Jy IL

sy i

_ OF THE

COUNCIL AND OFFICERS

ZOOLOGICAL SOCIETY OF LONDON.

TOTES

COUNCIL.

His Grace Tor Duke or Beprorp, K.G., President.

Tue Haru or Auramont, F.S.A.

Prof. J. Rosr Braprorp, M.D.,

D.Se., F.R.S., Vice-President.

Lt.-Col. Str R. HaAvetock-

Cuaruss, K.C.V.O., M.D.

AtrreD H. Cocks, Esq., M.A.

The Rt. Hon. the Haru oF -

Cromer, P.C., G.C.B.,

G.C.M.G.

F. D. Dawrrey Drewirt, Ese.,

M.A., M.D.

CuarLtes Drummonp, Ksq.,

Treasurer.

Sir Epwarp Duranp, Bt., C.B.

Freperick GILuert, Hsq., Vice-

President. -

Srpney F. Harmer, Hsq., M.A.,

Sc.D., F.R.S., Vice-President.

lb) Bs GOB.

| Srk Epmunp G. Lover, Bt.

Vice-President.

P. Cuoatmers MircHett, Esq.,

MIDAS. DSc), Hons WiLDy,

E.B.S., Secretary.

W. R. Ocinvin-Grant, Esq.

Aprian D. W. Potiock, Esq.

A. Trevor - Barrys, KEsq.,

M.A.

AntHony H. WHINGFIELD,

Esq.

A. SmirH Woopwarp, Esq.,

LL.D.,F.R.S., Vice-President.

| Henry Woopwarb,Ksq., LL.D.,

F.R.S., Vice-President.

PRINCIPAL OFFICERS.

P. Cuatmers MitcHeny, M.A

Secretary.

>» DSc. Hons iekess

Frank KE. Bepparp, M.A., F.R.S., Prosector.

ROWE Pocock, HRS. FAS:

Curator of Mammals, and

Resident Superintendent of the Gardens.

D. Sera-Smiru, Curator of Birds and Inspector of Works.

Henry G. Purmmer, F.R.S., M.R.C.S., Pathologist.

F. H. Warernouss, Librarian.

Joun Barrow, Accountant.

W. H. Cote, Chief Clerk,

Sir Warer Roper LAWRENCE,

E. G. B. Mrapre-WaA.po, Esq.,

<0 cA ete ali le sare

LIST OF CONTENTS.

1911, pp. 1-555

EXHIBITIONS AND NOTICES.

The Secrerary. Report on the Additions to the Society’s

Menagerie during the months of June, July, August,

eigjcimemollogie, ayaa! cursors IDI) op ccesgseoseeuoceodsoseds ax

Mr. R. I. Pocock, F.L.8., F.Z.8. Exhibition of a living

specimen of a Black Rat (dZus rattus) from Sark ......

Mr. R. I. Pocock, F.L.S., F.Z.8. Exhibition of a female

hybrid between a inale Black Lemur (Lemur macaco)

and a female of the Red-fronted variety of the Fulvous

Lemur (Lemur fulous rujifrons) ......0.0.c.c.ec es ecnc tenes

The Hon. N. Coarnes Roruscuinp, M.A., F.Z.S. Exhibition

of some Fleas, interesting in connection with the

Sobeadimonat PlAasMey. oct. vedas cask iacstnes cade aeeehen

Mr. J. Lewis Bonsots, M.A., F.L.S8., F.Z.8. 9 Exhibition

of, and remarks upon, a young Cairo Spiny Mouse

(Acomys cahirinus) about twelve hours old ...............

Mr, J. Lewis Bonuors, M.A., F.L.S., F.Z.8. Exhibition of,

and remarks upon, a pair of hybrids between the

Bramble-Finch (Fringilla montifringilla) and the

Chettinely (Hringillia celebs) ©. sexsccscs sc cccc sn sobs orinas senses

Mrs. R. Haic Tuomas, F.Z.S. Exhibition of a series of skins

illustrating an experiment in Pheasant-breeding, and

an account of the production of Thawmalea obscura in

the F, generation from a cross between 7. amhersti °

pied WAL EUGTEy dee Dandadcs dehuo see aoc Lone ny eee thee

Page

Dr. W. Nicott and Prof. EK. A. Mincuin, M.A., V.P.Z.S.

Exhibition of, and remarks upon, two species of

Cysticercoids from the Rat-flea (Ceratophyllus fasciatus).

(Text-figs: W052): cain wea ane Meee OEE aca ts.rcise =

Di. Wittiam Nicott. Demonstration of his method for the

collection of Trematodes

Dr. R. T. Lerprr, F.Z.S. Exhibition of photographs and

speclmens showing the Nematode infection known as

OnChOCerCIASIS) oF. 6 52 dss ae eee Rent Glee eee eee sraen

Dr. J. F. Gemuiin, M.A.. D.Sc. Notice of a paper on “The

Development of Solaster endeca Forbes ”

Mr. D. Sera-Suity, F.Z.S. Exhibition of, and remarks

upon, living examples of the Australian Budgerigar

or Undulated Grass- Parrakeet (I/elopsittacus undulatus),

showing three colour-phases

The Srcrerary. Report on the Additions to the Societys

Menagerie during the month of November 1910 ......

Dr. H. Hammonp SmirH, M.R.C.S., F.Z.S. Exhibition of,

and remarks upon, a specimen of the Red Grouse

(Lagopus scoticus) which displayed a curious variety

Chard NR OV MONTE HAA | OMUNTEVELS 2 0, J os20ndpqadacoesbonsenbacnaocnsn

Mr. D. Sera-Smiru, F.Z.S. Exhibition of some skins of the

Australian Yellow-rumped Finch (Munia flaviprymna)

showing markings tending towards those of JM. castanet-

BROT OE (55 Jo oe SLE SAORSE SEAT SER REE EEE RERY AL BERET eee

Dr. W. E. Hoyts, M.A., F.Z.8. Explanation of the Report

of the International Commission on Zoological Nomen-

clature presented to the Graz Meeting of the

International Zoological Congress

The Sucretary. Report on the Additions to the Society’s

Menagerie during the month of December 1910 ......

Dr. G. W. Anprews, F.R.S., F.Z.S. Exhibition of the

skull of a Sabre-toothed Tiger (Similodon califorinicus),

from an asphalt deposit in California .....................

The Secretary. Exhibition of a mounted specimen of the

Platypus (Ornithorhynchus paradoxus) ...116...eceeeeee

Page

100

101

181

Mr. Epwarp GerrarD, F.Z.8. Exhibition of the head of a

Caribou (Rangifer tarandus carihow) with abnormal

antlers. Also the heads of an Eland (Zaurotragus

derbianus gigas), and of three White-tailed Deer (Dorc-

elaphus americanus macrurus), (Text-fig. 52) .........

The Szcrerary. Report on the Additions to the Society’s

Menagerie during the month of January 1911 .........

Dr. H. Hammonp Smiry, M.R.C.S8., F.Z.8. Exhibition of

three skins of male Pheasants assuming female plumage

Mr. G. A. Bounencer, F.R.S., V.P.Z.8. Notice of a paper

“On a Collection of Fishes from the Lake Ngami

Rasim, Iacovella’ ccédoccs aceneosesocded concveseodeneeae

Mr. Aurrep H. Cocks, M.A., F.Z.S. Exhibition of photo-

graphs of, and remarks upon, the female Brindled Gnu

recently born in the Society’s Gardens. (Text-figs. 97

Se GIS) y cx culbspa bone shi seahunmeabaupeacose au asa cemeenpuEaHBecmaaas

The Srcrerary. Exhibition of a series of lantern-slides

illustrating some of the most remarkable features of

Mr. Carl Hagenbeck’s new Tierpark at Stellingen ......

Mr. Ernest C. OperHoirzer. Exhibition of lantern-slides

and photographs illustrating an account of some

(QOSeRATaNELONNS) Cray I Kole Nodhe coobosonadacrocbusonoecnoduacpeac

Page

181

314

308

Ne)

PAPERS.

. On the Inheritance of the Webfoot Character in Pigeons.

By J. Lewis Bonuorz, M.A., F.LS., F.Z.8. (Text-

IMSS) Gas 445) seopdacnesodsosoroh bodes Scogaubagsncovsneq0680080a000

Notes on the little-known Lizard Lacerta jacksoni Blegr.,

with Special Reference to its Cranial Characters. By

Epwarp Dscen, F.Z.8. (Text-figs. 5-7.) ...............

On the Peloponnesian Lizard (Lacerta peloponnesiaca

Bibr.). By G. A. Bovutmncer’ F.R.S., V.P.ZS.

(Plate I. and Text-fig. 8.)

eee cee sere ener reese esereses esse sees

. Remarks on Two Species of Fishes of the Genus Cobdius,

from Observations made at Roscoff. By Hpwarp G.

BouLENGER

eee cece r reco ee sors eee ees eseoeseeooseyooessososseessane

. On a Possible Cause of Pneumo-enteritis in the Red

Grouse (Lagopus scoticus). By H. B. Fantuam, D.8c.,

B.A., F.Z.S., and H. Hammonp Sire, M.R.C.S.,

i Dre 53 Ol eae Cie) Zh wo aR a0 5.40 Ma Nout ogSeokSosseHuoddaddoanae

On the Alimentary Tract of certain Birds and on the

Mesenteric Relations of the Intestinal Loops. By

Frank E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to

thie Socieby. ) (extaties 19 =28)) ee aeeesee ance ene eetieee

On the Specimens of Spotted Hyznas in the British

Museum (Natural History). By Prof. ANGEL CABRERA,

ORIN IAs Bee Me oa AU RS GL ARAL Ah ROR GOs 80 60:

On the Segmentation of the Occipital Region of the Head

in the Batrachia Urodela. By Epwin 8. Goopricu,

M.A., F.R.S., F.Z.8., Fellow of Merton College, Oxford.

(M@ext=figs: 29-51.) coe. ise sts etic hictes's « cieitce Meee eee ETE

The Mammals of the Tenth Edition of Linnexus; an

Attempt to fix the Types of the Genera and the exact

Bases and Localities of the Species. By OLpFriEeLp

MR roy eee IM] bhiob lle Ade Pre cot onan en annongktednoabasutact daa:

Page

101

120

10.

13.

14.

15.

16.

ESE

19.

20.

Vil

The Duke of Bedford's Zoological Exploration of Eastern

Asia.—XIII. On Mammals from the Provinces of

Kan-su and Sze-chwan, Western China. By OLprreLp

AIDETOMUAS VME EUS See RL 15S ep jac. ecciensais-s oyee-c.ave ayeisis vie ed Re

On the Structure and Function of the Gas Glands and

Retia Mirabilia associated with the Gas Bladder of

some Teleostean Fishes, with Notes on the Teleost

Pancreas. By W. N. F. Woopiann, F.Z.8., The

Zoological Department, University College, London.

(Plates II.-IX. and Text-figs. 53-62.) ............cce eee

. On Skulls of Oxen from the Roman Military Station at

Newstead, Melrose. By J. C. Ewart, M.D., F.BS.,

EAS aen) (Nextatio ses Os Ol) mu remrret conte ls aseasos esol. ve

Plankton from Christmas Island, Indian Ocean.—l. On

Copepoda of the Family Coryceide. By GrorcE P.

IGPACURY Nome (ra: bes ONG NOU Vibe oe naka ccgne marae a. sea. 2

On some New Zealand Spiders. By H. R. Hoge, M.A.,

Ee Ze S em exci tig sty 2-008) Wy see caree secs aie acs seine

Report on the Deaths which occurred in the Zoological

Gardens during 1910. By H. G. Primer, F.RS.,

‘F.Z.S., Pres. R.M.S8., Pathologist to the Society.........

A Contribution to the Study of the Variations of the

Spotted Salamander (Salamandra maculosa). By

Hpwarp G. BouLencer. (Plate XV. and Text-

(igay” Cz Ry UA SNE ais ce untae Pe eeaege oo

On the Mountain Nyala, Tragelaphus buxtoni. By R.

LypEKKER. (Plate XVI. and Text-fig. 103.) ............ é

Observations on different Gibbons of the Genus Hylobates

now or recently living in the Society’s Gardens, and on

Symphalangus syndactylus, with Notes on Skins ijn the

Natural History Museum, 8. Kensington. By Dr. F.

STDS VAHL CI Zit Septet eset ny eeces tre gee of Uke cats te ie shai Se oo é

Some New Siphonaptera from China. By Kart Jorpan,

Ph.D., F.E.S., and the Hon. N. Cuartes Roru-

scHILD, M.A., F.Z.S., F.E.S. (Text-figs. 104-124.) ...

Contributions to the Anatomy of the Anura. By Frank

E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the

Society. (Text-figs. 125-133.) ...0.....ccceceeeeeeeeeeeee nes

Page

158

249

318

369

393

Wiil

21. Gn the Spermatophores in Harthworms of the Genus

Pheretima (= Pericheta). By Frank E. Bepparp, M.A.,

HARSs) Rez: ao Prosector to the Society. (Text-figs.

134- 136.) ieee BP hy Sa pa AP IMIR eee

22. A Rare Beaked Whale. By R. Lypexxer. (Text-figs.

UY ee eo) Yastates a. AMG MERA ASS has LSAT POR ok ics ORR BO or

23. Age-Phases of the Rorqual. By R. LypExKER

24. On Longevity and Relative Viability in Mammals and

Birds ; with a Note on the Theory of Longevity. By

P. Coatmers Mircnent, M.A., D.Se., LL.D., F.R.S.,

Secretary to the Society

Cenc ee mee meres ee pere erro ss eresseceeane

25. Some new Parasitic Nematodes from Tropical Africa.

By Roserr T. Leper, D.Sc., M.B., F.Z.S. (Text-figs.

MAO AAA. J ish od te eoses aie laf cia te ah oe MRR iN ee

ADDENDUM.

A correction to Dr. R. T. Lerpmr’s Memoir on the “ Entozoa

of the Hippopotamus ” 3

i On Sn ii aC aan iki a ke cr a i ney

Page

425

549

AV ALIP inl Je) OMe CAN 1) ty

OF TIIK

CONTRIBUTORS,

With References to the several Articles contributed by each.

(1911, pp. 1-555.)

Page

ANDREWS, CHARLES WiuLiAM, B.A., D.Sce., F.R.S., F.Z.S.

Exhibition of the skull of a Sabre-toothed Tiger

(Smilodon californicus), from an asphalt deposit in Cali-

IRODALIUIEL cies esa aah al Mle eae Ata po Ae aa an 181

BEDDARD, Frank E., M.A., F.R.S., F.Z.S., Prosector to the

Society.

On the Alimentary Tract of certain Birds and on the

Mesenteric Relations of the Intestinal Loops. (Text-

HSM 1S) Bas Mae SeE ERT Tae | sree Eid raeeine sc cise eises/s scion eoeee 47

Contributions to the Anatomy of the Anura. (Text-

HES, WAD=N238)5)): coaee see o0ce pene conanesandeqekeceboE UdbaeeEHEHOee repr 393

On the Spermatophores in Earthworms of the Genus

Pheretima (=VPericheta). (Text-figs. 134-136.)............ 412

Bonnore, J. Lewis, M.A., F.LS., F.Z.S8.

Exhibition of, and remarks upon, a, young Cairo Spiny

Mouse (Acomys cahirinus) about twelve hours old .........

Exhibition of, and remarks upon, a pair of hybrids

between the Bramble-Finch (fringilla montifringilla) and

the Chattinels@h7nqille: calebs)\\. <).ta.css-o2 rset tac s-

On the Inheritance of the Webfoot Character in

Pigeons cum @bemtenOs.y3 65 4.) cin /scon enlace nee eeeeGreeen cnet

BouLencer, Epwarb G.

Remarks on Two Species of Fishes of the Genus

Gobius, from Observations made at Roscoff ..................

A Contribution to the Study of the Variations of the

Spotted Salamander (Salamandra maculosa). (Plate XV.

and Text-tigs. 99= N02.) - ei len sine cn eere eee eee seer ne ere

Bou.encer, Grorce A., F.R.S., V.P.Z.S.

On the Peloponnesian Lizard (Lacerta peloponnesiaca

Bibrs)m ((@elatediiamd Mlext tow St) ereyece eee sapere ertttat:

Notice of a paper ‘Ona Collection of Fishes from the

Wake Neami ‘Basin, Bechuanaland yeas epece es seeee ee

Caprera, Prof. AncEL, C.M.Z.S.

On the Specimens of Spotted Hyznas in the British

Museum (Natural ieastory) i 20-0242. 4... anced eee a eeeeeee

Cocks, ALFRED H., M.A., F.Z.S.

Exhibition of photographs of, and remarks upon, the

female Brindled Gnu recently born in the Society’s

Gardens. -(Mext-figs: O77 i 98.) Me... ndeesemaee cacti oh

Page

323

314

DrGEN, Epwanrpb, F.Z.S.

Notes on the little-known Lizard Lacertajacksoni Bler.,

with Special Reference to its Cranial Characters. (Text-

INES, D=(/ 0) cass Alomhe cea nee HARE RH aT te an eenernEe eiea ale

Ewart, James Cossar, M.D., F.R.S., F.Z.8S.

On Skulls of Oxen from the Roman Military Station

at Newstead, Melrose. (Text-figs. 63-91.)..................

FantHamM, Haroup B., D.Sc., B.A., F.Z.8S., and Smrru,

H. Hammonp, M.R.CS., L.R.C.P., F.Z:8.

On a Possible Cause of Pneumo-enteritis in the Red

Crousey CLAgGOUS SCOLLCUS) a0. na eco aaa aie cnacsdcccd so dedas

Farran, Grorce P.

Plankton from Christmas Island, Indian Ocean.—lI.

On Copepoda of the Family Coryceide. (Plates X.—XIV.)

GemMILL, Dr. James F., M.A., D.Sc., F.Z.S., Lecturer on

Embryology in the University of Glasgow.

Notice of a paper on “The Development of Solaster

CRCCEMB OL DES) Mh Aoi npiants dante so eis < thiaipiie Sissies te teiseSes sese ee ao

GERRARD, Epwarp, F.Z.S.

Exhibition of the head of a Caribou (Rangifer tarandus

caribou) with abnormal antlers. Also the heads of an

Eland (Zaurotragus derbianus gigas), and of three White-

tailed Deer (Dorcelaphus americanus macrurus). (Text-

1G, DAD I BAn eit drsch cbt dbo SHe pee bonebe bo, CRORE CEE Mer a aan eee

GoopricH, Epwin §., M.A., F.R.S., F.Z.S., Fellow of

Merton College, Oxford.

On the Segmentation of the Occipital Region of the

Head in the Batrachia Urodela, (Text-figs. 29-51.)

Page

249

282

181

101

Page

Hoge, Henry R., M.A., F.Z.S.

On some New Zealand Spiders. (Text-figs. 92-96.) ... 297

Hoyts, Dr. Witi1aM E., M.A., F.Z.S., English Member of

the International Commission on Zoological Nomen-

clature.

Explanation of the Report of the International Com-

mission on Zoological Nomenclature presented to the

Graz Meeting of the International Zoological Congress... 101

JorDAN, Karu, Ph.D., F.E.S., and Roruscuiip, The Hon.

N. Coaruss, M.A., F.Z.8., F.E.S.

Some New Siphonaptera from China. (Text-figs.

1) CRY) Aa fae eared ado aGobn oko adotoS en sontnass5o0¢ 365

Leirer, Rosert T., D.Sc., M.B., F.Z.8.

Exhibition of photographs and specimens showing the

Nematode infection known as Onchocerciasis ............... Ad

Some new Parasitic Nematodes from Tropical Africa.

(berg mils WHOS) pocoecasnonsdo02cnnacaccoasnsageannscoase 0F¢ 549

A correction to the Memoir on the ‘ Entozoa of the

JBI oy s10) OWE NETO © Ghunonansoa does asocdsanosocsadoasdeonasuacsn9oqne0s 555

LYDEKKER, RICHARD.

On the Mountain Nyala, 7'ragelaphus buxtoni. (Plate

XV 1. and Vextafie slOds)\ (ic 5 k,l een emotes sls eae yanare 348

A Rare Beaked Whale. (Text-figs. 137-139.) ......... 420

Ager hasesiol, he shorqualll sede... 5- see eee 423

Mincurn, Prof. Epwarp A., M.A., V.P.Z.S., and Nicott,

Dr. WitiiaM, F.Z.S.

Exhibition of, and remarks upon, two species of Cysti-

cercoids from the Rat-flea (Ceratophyllus fasciatus).

Wepre itersy I Gs 5) aan nagedoctinnwes. qosmdapencconacu soda jaqsonne” 9

xil

MircHetyt, P. Cuatmers, M.A., D.Sc., Hon. LL.D., F.R.S.,

F.Z.8., Secretary to the Society.

Report on the Additions to the Society’s Menagerie

during the months of June, July, August, September,

and October, 1910

CO a a Cn a SO i ei i ie i i i ar ad

Report on the Additions to the Society’s Menagerie

during the month of November 1910

Peewee cee ese aes eee as aene

Report on the Additions to the Society’s Menagerie

during the month of December 1910

Ce i

Exhibition of a mounted specimen of the Platypus

(Ornithorhymehus paradowus) 0. s2o..2.c2- 2: escuns essed dees 2s:

Report on the Additions to the Society’s Menagerie

during the month of January 1911

Exhibition of a series of lantern-slides illustrating

some of the most remarkable features of Mr. Carl

' Hagenbeck’s new Tierpark at Stellingen

On Longevity and Relative Viability in Mammals and

Birds ; with a Note on the Theory of Longevity

Nicouz, Dr. WixiiaM, F.Z.S., Lister Institute of Preventive

Medicine.

Demonstration of his method for the collection of

Trematodes

i i i ii i ei i ink i ii i ei iii i ee ee ee iy

Nicotz, Dr. Wiii1am, F.Z.S., and Mincuin, Prof. E. A.,

M.A., V.P.Z.S.

Exhibition of, and remarks upon, two species of Cysti-

cercoids from the Rat-flea (Ceratophyllus fasciatus).

(Text-figs. 1 & 2.)

Ci i i ei i i i i i ii acacia

Page

100

181

18}

314

358

425

X1V

OBERHOLTZER, ERNEST C.

Exhibition of lantern-slides and photograph, illus-

trating an account of some Observations on Moose ......

Purmer, Henry G., F.R.S., F.Z.8., Pres.R.M.S., Patho-

logist to the Society.

Report on the Deaths which occurred in the Zoological

Cardensralummme: GOO Re crea ce eee e ee oeeeree Neavcorsan et

Pocock, Reernatp I., F.R.S., F.L.S., F.Z.S., Curator of

Mammals, and Resident Superintendent of the

Gardens.

Exhibition of a living specimen of a Black Rat (J/ws

TRG AATES) oe ROM OPN Ce RESE en Dano adun Sous jn oocobhoabssnocesceds oc

Exhibition of a female hybrid between a male Black

Lemur (Lemur macaco) and a female of the Red-fronted

variety of the Fulvous Lemur (Lemur fulvus rufifrons)...

Roruscurip, The Hon. N. Cuaruss, M.A., F.Z.8., F.E.S.

Exhibition of some Fleas interesting in connection

THIGH OE ep OTOANOUUENE? CVE THEY) “bon < dee sosegeacsonousagusedoouoce

Rorsscuitp, The Hon. N. CHaruss, M.A., F.Z.8., F.E.S.,

and JoRDAN, Kart, Ph.D., F.E.S.

Some New Siphonaptera from China. (Text-figs. 104—

MS VAN) ES ROS haat BG DERG ee eel bese ok heute s eae ee

Sers-Smira, D., F.Z.S., Curator of Birds.

Exhibition of, and remarks upon, living examples of

the Australian Budgerigar or Undulated Grass-Parrakeet

‘(Melopsitiacus undulatus), showing three colour-phases ...

398

318

365

Seru-Surra, D., F.Z.8. (Continued.)

Exhibition of some skins of the Australian Yellow-

rumped Finch (dunia flaviprymna) showing markings

tending towards those of WM. castaneithoraa

ee ae

Smira, H. Hammonp, M.R.CS., L.R.C.P., F.Z.8.

Exhibition of, and remarks upon, a specimen of the

Red Grouse (Lagopus scoticus) which displayed a curious

Vabiety Ol the ORGIMarye PUIMAMe! ee cc.- 0: on-cre es s2c+e <n ss

Exhibition of three skins of male Pheasants assuming

eatin © OMIM OR ese es toay-tacyies cecil cin sider SATS cin 2 = tar ce as os

Smira, H. Hammonp, M.R.C.8., L.R.C.P., F.Z.8., and

Fantuam, Harovp B., D.Sc., B.A., F.Z.S.

On a Possible Cause of Pneumo-enteritis in the Red

Grouses(Zagopie SCOUCIS)) Cs... ack ontc oe eta ne seanc esos

Tuomas, OLDFIELD, F.R.S., F.Z.8.

The Mammals of the Tenth Edition of Linnzus; an

Attempt to fix the Types of the Genera and the exact

Bases and Localities of the Species ...................0.2000

The Duke of Bedford’s Zoological Exploration of

Eastern Asia.—XIII. On Mammals from the Provinces

of Kan-su and Sze-chwan, Western China ................0.

Tuomas, Mrs. R. Hare, F.Z.S.

Exhibition of a series of skins illustrating an experi-

ment in Pheasant-breeding, and an account of the

production of Thawmalea obscura in the F,, generation

from a cross between 7’. amhersti 2 and 7’. picta 3 ...+.

101

100

314

xvl

t Page

Wetcu, Dr. Freperick D., F.Z.8.

Observations on different Gibbons of the Genus Hylo-

bates now or recently living in the Socicty’s Gardens, and

on Symphalangus syndactylus, with Notes on Skins in

the Natural History Museum, S. Kensington ............ 393

Wooptanp, Wittiam N. F., F.Z.8., The Zoological Depart-

ment, University College, London,

On the Structure and Function of the Gas Glands and

Retia Mirabilia associated with the Gas Bladder of some

Teleostean Fishes, with Notes on the Teleost Pancreas.

(alates Me TXevand Mext t1oS5/93—022)\epeceee nena. cer are 183

XVil

LIST OF PLATES.

1911, pp. 1-555.

Plate Page

eEacenianpelaponnesiace: Whig caine halt oe en ee es 3

ut a

Wee GastGlandsvoth Meleastenemts vee 6. a. ee ona oe ee eee 183

vil |

ee 1=6: Corycella brews: 7-VN.- OL curte-.25:9..2 oe 5

XI. 1-6. Corycella curta, 7. C. brevis. 8,9. Coryeella sp. 3.

10. C. earinata. 11,12. Coryceus gracilicaudatus.

XII. 1-8. Coryceus cutus. 4-6. C. obtusus. 7. C. dubius. ye

Si Gis Ca CANS Ferra ces yRy ra Aare te AYR a she Ai Rca FAVS Ay gaat

ROI 16s Coryeeus murraye 7-9. Ci andrewst. van ccielseus »

XIV. 1-4. Coryceus andrewst, 5-9. C. dubtus .......000050, !

XV. Salamandra maculosa var. teniata, var. mollert, and forma

CY PIUC i. «a as alleen oli ite Aa teat epee Wes P< cei old rc 325

XVI. The Mountain Nyala, Trageluphus burtont ....... 0.0.45. 348

Proc. Zoon,. Soc.—1911,

LS) I

ho bo bo bo to bt

OUR OO to

SP)

XVill

LIST OF TEXT-FIGURES.

5 (Cy Saleareconel Ole EO MeCN DOS UMA 45504069055; sandeoeaes

Pa ysnlcercordsolela/menvole pes 210000.) tek eve ele nena ae

peBedigree of Minn Smalley:s pigeon (W/69)) 9) eee sae

. Genealogical Table showing matings and results in mixed strains

OU PME CONS Siete cv eelenineret, Sinan MEO ROR Sty. eee

. Upper, lower, lateral and posterior aspects of skull of male

WigcentaqUchsont Fey oc\ Hee ere ee eee erate

. Lamina supraciliaris and supraorbital bone of Lacerta jacksone.

Also showing configuration of nasal process of premavillary

and outer nasal apertures of L. jackson, 3, 2 3 L. muralis,

i éypica, 6; 1. veipara,) 6; anda, agilisi eric: see

. Upper, lower, lateral and posterior aspects of skull of female

LEGCOF LET OCESONT here tan eM ete eee CER ECE Oa eee Oe

. Upper and side views of head. of male and female Lacerta

CLOMONTICSTA COI BEk ek eee nos ee EE ee eae

Rulntestinall tract olerynemnanuenican@mr i) iw selon eee aie

NalintestanallitractioteCyy bene Succi tac avert rte eye) ten race

elmtestimalitrachiotNalegal an laccriie mee wetter tener ee

intestinal tract otiOntalsmujicauda ae eri ere

. Intestinal tract of Huplocamus nycthemerus, showing condition

TRONTCVENG| INROVEN TNS) MOMTIAl, oh canonbooaronocsasdpH no donne

elmitestinelletractioneloa anowsiCWOrci. a. eeris etree tnt 1 eer ene

> intestinal tract of Gymnorhina lewconota “ein aie cl.) ele. te

Intestinal tract of Melanerpes supercitiaris ......e.eee eres eee

_ liars! HACE Ot GHAGIANIS BORIS 5 200cnncacceoosnnobonencor

5 Jnana eRtee OF CRS JODORUGIS <556n556000500008505525500

| Thatiegniiaall weanets © Wowie BONG 460600000092 040s0v0000000"

Palintestinalutractio Sprzaerus Well Costs s.. - leis teen neneetets

—

Thnibectinaaul ees Git OMiMAGHRIZS CHEE 5460504c000000es5000n 45 0ne

TionRaciauaal HRA Le 41510 DUIS 5rbnoonncagedocdocccoudaasvosans

salmtestinalemacthiol eipodotusaustiialis: ln ott ed eee aerate

. Intestinalitrachioh Biuvanus equips .- = 22 eee eee oe

. Intestinal tract of Gtdicnemus scolopat «1... eee ee cee ee eee

, Imtestinalitiract on hnatencula aicivea hi. + see ene eee

X1x

Paga

27. Intestinal tract of Tachybaptes fluviatilis oo. vec cee cee oll

98. Intestinal tract of Nestor notabilis ............: eis uubns BAG 84

29, Dissection of a full-grown Axolotl (Amdblystoma tigrinum),

showing the skeleton and nerves of the head and three trunk-

SEPANG IU Smistered saris: amettiafate: Pe yee o's) s) ae: ale ale et sane che pee aterm 104

30. Outline of spinal cord and hind brain, with roots of cranial

nerves and first two spinal nerves, of a full-grown Axolotl.. 108

31, 82, Partial reconstructions of the hind region of the skull and

the anterior region of the trunk of an advanced larva of the

PNSCONGH We ios ee he Aewhn birch clo ec His Cen oa ae 105, 106

33-30. Partial reconstructions of the left side of the head region of

Aiyouney lar venom Chem col Oblitiay. sale este aslels se). -) on LOG OT

36. Reconstruction, seen from above, of the hind region of the skull,

and anterior neural arches of a young larva of the Axolotl .. 107

37. Left side view of a partial reconstruction of the anterior end of

the trunk-region of a young larva of the Axolotl.......... 108

38, 39. Partial reconstructions, seen from the right side, of the

anterior end of the trunk-region of a young larva of the

PACK OL OU tarts carn Severe ohana net eeene Ved Lae eee ae 108

40. Reconstruction, seen from above, of the head and anterior |

trunk-region of a young larva of the Axolotl.............. 109

41, 42. Reconstructions of the anterior trunk-region, seen from the

right side, of a young larva of the Axolotl ............ an LO

45. Left side view of a reconstruction of the anterior trunk-region

OigapyounovlanrvarOl the 7A COOL UNE. aes we a/c Cites dite 57s lo iBaL

44. Right side view of a reconstruction of the anterior end of a

LeEyan Oils ie NeOlOULLN: sracteeice.aatienten anh ua went stale cutee, ti}

45, Partial reconstruction, seen from above, of the anterior region of

cinwe rn yO) OlabhewAmcOlat lava temient te « 1c tetra aetaeaey pets 112

46. Similar reconstruction of the more dorsal region of the same

IASKG lot cma DT VAOW ays cette che «F.C ume a eeoke er aay ela 113

47. Left side view of a reconstruction of the anterior region of an

embry orAscolobl ty acer. «wis s vaenc ie oe Pe ee wag 113

48. Partial reconstruction of the anterior region of an embryo

Axolotl seen from abaven ay tp cta emis hen see eine cha raters 114

49, Right side view of a reconstruction of an embryo Axolotl .... 115

50. Diagrams of the segmentation of the metaotic region in

WENTOH OY NU) OVE Rotate enews cece SIGE ia brie Roe iene aa ee 116

51. Diagram of the segmentation of the anterior region of an

“LounpliciiRern PT Ande iy Pity he eco Ont I Ie 116

52. Head of a Caribou (Rangifer tarandus_caribou) ith abnormal

SUINGLETSM | 5, 40. ot oy Aer MR RY MVR: Lely sLptouas Auch sh chslo ce Se odal | wh aiah a 182

53. The bladder, duct, and “ red bodies ” of Angzlla vulgaris, viewed

SM OVETI TTR TSTO RE ics. tle ee eee om CL eae Ve 186

b4: External aspect of an Hel’s rete mirabile 0. .......0.. 000-00 u 187

55. The construction of an Kel’s rete mirabile bipolare geminum .., 188

XxX

Page

56. Vein surrounded by modified acini of the pancreas in Nerophis

EQUOPUUS Ne ic tate IO EC EE BE TA Ree 199

57. The construction of the “ved body ” of Peristethus cataphractus . 202

58. Diagram to explain the probable construction of the gas glands

of Trigla gurnardus and Smarts maurtt . 0... ccc. cece ee eee 207

59. Diagrammatic transverse sections through the anterior end of

“red body ” in ventral wall of bladder of Cepola rubescens .. 213

60. Vacuoles in the kidney cells of Nerophis ........-.0000cceuee 225

61. Microphotograph of transverse section across thin region of gas

gland of Perca, showing foam-like mass of bubbles on surface. 240

62. Microphotograph of transverse section across thick region of gas

gland of Perca, showing extruded gas bubbles on surface.

The “spent” condition of the cells is also shown ........ .. 240

G3 eSicullliotthe Anos (Bos! depressicon7s)\ne seen ele eee ee 250

64, Front part of the skull of a Buffalo (Bos bubalus) ............ 251

65. Front part of skull of the Urus ( Bos primigenius) ............ 951

66. Skull of a Urus in the Anatomical Museum, University of

ASUisallotumpsdn Gee ive, ante ©: aes aretha ne Oe ee Mies ote de oe 252

Git ront part ot siculljof American! bisonlepaer eee eee 253

68. Front and side views of the skull of a young fetal Ox ........ 258

69. Front view of a foetal Ox skull about the fifth month, and front

partjof same shitlliy cs cL pi emia es ieee eRe ee oe 255

70. Front view of Calf’s skull at birth, and front part of same skull. 257

71. Front part of skull of a polled Aberdeen-Angus Ox .......... 258

72. Skull of a Syrian Ox with vestigial horn-cores and a forehead

like that of flat-polled Aberdeen-Angus cattle ............ 259

73. Front part of skull of polled Newstead Ox in which the upper

end of the premaxille biftircates .............. Sen ee ame eme 261

74, Front part of a horned Newstead skull in which the premaxillee

are short and fail to reach the nasals ............ se HIS 262

76. Front part of skull of a horned Cadzow Ox with notched pre-

maxillee which fail to reach the nasals .................. 263

76. Front part of skull of a cross-bred Shorthorn in which the pre-

maxille fail toaeach thie masalsiey.. © reaciscies celia welt 263

77. Skull of polled Newstead Ox of the type, which, with horns, are

sometimes said to belong to Bos frontosus Nilsson .......... 264

78. Right half of occiput of the Aberdeen-Angus skull with long

MmaMmedlbss MNO A SXCMIOGM OEMS ~Gpoesouccooscgosacce 265

79. Outlines of the occiput and part of the forehead of three Urus

ISU UIST lc AG rie er ree era Ieee al ona RES SCI, G..6.d.cuh!o, Hin. ola 266

80. Front view of skull of the Bison (Bos biven) ............-05- 267

81. Hind part of skull of the Urus represented in text-fig. 66...... 268

82. Frontlet and horn-cores of a small Newstead Ox with a

promimentaioneleadd yr. stacy 00 - «ce etter teem tie ee 269

83. Occiput of the Urus skull represented in text-figs. 66, 79, and 81. 270

87. Front view of skull of a white “wild” Cadzow Ox from

Hamilton Park. Also occiput and horn-cores of same

sill SoS 5550559505 AS AS 3.cgn cca, 0 ERC aR ER ee Re 276

88. Occiput and horn-cores of an American Bison .............. 276

89. Front part of the skull of a Zebu (Bos indicus) in which the

TOWEINE NAMES THER ATE TRISENISH 3 on 6n nia oe 5 adn MOS ano 278

90. Front part of the skull of a Zebu in which the premaxille fail

homenehmulernedselswewy.eilee, Ps jemi e erie pean cee erases ece ome 8 279

91. Occiput of the Zebu with the premaxille reaching the nasals,. 280

a LICLRAGNALROMICTOL Valin ChOSSI at NIGLOWm tars Gite siejrelere 24.04 <4. «1 301

Sam GHOnN aires miemaley Warner de. tine ne: Meise en iela ae «+ 304

UApecncoctenusnaurcuses Hemalevs x. acserten ot ee eee vse creo teas 306

SERUM MOGICRESRE RULE GTID A thes felinfuilaii Pose faite lel Elke Tree ine iis. «i's Miavare si 509

SemeYolomedesriinentagus., «Kemala racine ae ae cen oa ti 311

97. Young Brindled Gnu born in the Society’s Gardens.......... 315

OS mileadsoyoune irindled (Gmury sos. cere se aielets ese a « 316

99. Diagram to explain the tables of different markings in Sala-

TUITE, TACOS or a Dio) 6b oboe Ob OND Db Cob on UNS heer 025

1C0, 101. Salamandra maculosa forma typica...........6005- 329, 352

O02 Salamandia maculosa vans tcenidtu ..o. Seo... ce se cece sae 039

103. Head of an old Buck of the Mountain Nyala................ 351

104. Clasping organs of Archeopsylla sinensis G .........2+ eee 366

105. Abdominal segments VIT & VIII and zecepeaenl un seminis of

PACH CO PSH UGH SUNCNSUS Die acy Ale vate Sa Ae cle etr ele) oeeas sche 366

HOG aleadvon Ceratopheytlustcnmapus! Goss se nee ens eee en 367

107. Clasping organs of Ceratophyllus crispus $ «1... s eee eee 310

108. Abdominal segments VII & VIII, stylet and receptaculum

scummisrol Ceratophylims crispus DIN... ws cece tenes: 371

169. Abdominal segments VII & VIII and stylet of Ceratophyllus

COLGCTISNODY ON RT ae ON ee are nae te ea tee ees 371

110. Clasping organs of Ceratoph ples CANNES DOUS So ech OD DDC DOS ON 373

111. Abdominal segment VIII and receptaculuin seminis of Cerato-

DY LLUS CLASSUSE OMG AES SAY RAL cote eee Ne ote este a cos. as Yi

112. Clasping organs of Ceratophyllus mandarinus G .........+4. 317

115. Abdominal segments VII & VIII and receptaculum seminis of

COR MLO Phy lUS ORBITS OM esas sen chslaye ae ee ns a ee 378

114. Clasping organs of Ceratophyllus mongolicus S ....... 0000005 379

115. Abdominal eomhane VIL & VIII ard receptaculum seminis of

XX1

Page

. Occiput of the Newstead skull represented in text-fie. 86 .... 271

. Occiput and horn-cores of a Urus in which the notches below

the horn-cores are deeper than in text-fic.83.............. 272

. Front view of the Newstead skull of which the occiput is

represented in text-fig. 84. Also the temporal fossa of same

shill <3 cs 5seouenoses Polshsvopsdararstote slovi cid a ce lerst ete Sete eee 274

(CERO OUI OL UBUITIS:, Osu Os sto aR RICE EB 381

121. Clasping organs of Neopsylla aliena 3

122, Abdominal segments VII & VIII and receptaculum seminis of

NeOp SIGUA » aa: Naya nade avo Uk ee eS 390

12a Eleadtotestenaponia coclestis) Os nnn rset eee 392

124, Abdominal segments VII & VIII of Stenoponia celestis Q .... 392

125. Ventral view of anterior part of the body of Megalophrys fee

Pabcralliy G1SSeCted -,..: ics iotalnie,h <ekee ete aGkoree ee nee 394

126. Sternum of Megalophrys fee from the ventral surface ........ 395

127. The alimentary tract of Megalophrys fee from the middle of

the stomach to the middle of the colon laid open and divided

Ito LWOPORWONS: 1... aaah nee La ence EAR ee 398

12S Gular sac oneMicgalophiysyicce sre see AOL

129. Ventral musculature of the throat of Megalophrys fee........ 403

130. Alimentary tract of Breviceps gibbosus with the greater part of

the smvallllimitestimeslaidwopen er eee en eine eee 407

181. Breviceps gibbosus, from the dorsal surface, partly dissected .. 409

132. An enlarged view of a portion of text-fig. 151, to show absence

of diverticulum of bedy-cavity overlying thigh ............ 410

133. Certain parts in the anatomy of Breviceps sp. .............. 41]

154. Spermatheca of Pheretima montana containing spermatophores. 414

135. Longitudinal section through spermatophore of Pheretima

IMONEANG. len hirer Tae Ee Ye Cho ebateaey Ee Ee aca: 416

136, Transverse section through wall of spermatophore of Pheretima

UAL aera Sman IN SR OS BAO OOO Cb ie Am Rew ciate G OF CO): 417

187. Mesoplodon (Disuiea en) GY UYU Pts Ae). Wide Nene R eae AONE SU te 420

138. Palatal aspects of cranium and lower jaw of Meéeseplodon

(DWiopledon) G7 Gye cya. ein ae eee bi eee Ten ee eee 421

139. Lateral view of skull of Mesoplodon (Dioplodon) grayt ...... 422

NAD, Valo CANO KOTO TU HISODE Sodde osoctooBs0ddauabnecoabagoc6s 550

NA eCylindropharyna Onevicdude pe y)n tt. eo ree eee 552

142. Cylindropharynex longicauda ... 06. cee eee eee oa pe ROR 508

LASS i aChYPhRary Nk RAGEKUC onc ce eas). 0+) te ate ete 554

WAAS ISPUODECEA UGANAG. ccayere Dalat a tinial. ths dhe ee ES 505

Page

. Abdominal segments VII & VIII and receptaculum seminis of

Cer atop hy llus euicles sO teers Reh Leer eee oe ee 383

7. Abdominal segments VII & VIII of Ceratophyllus pheopis .. 383

. Clasping organs of Amphipsylla casis G......--..--05----- 386

. Abdominal segments VII & VIII and receptaculum seminis of

Amphipsylla Casts sO wena) cg Acer eae ee us oe See 386

. Abdominal segments VII & VIII and receptaculum seminis of

IN COp Sy Wa COMBO? Cited... tyke. alee eh, eee Mee ROR 387

xxii

NEW GENERIC TERMS

PROPOSED IN THE PRESENT VOLUME (pp. 1-555).

Page Page

Acheilostoma (Vermidea) ...... 549 Cylindropharynx (Vermidea) ... 551

Cobboldina (Vermidea) ......... 505 Stenoponia (Siphonaptera) ...... 391

Corycella (Crustacea) ............ 283 Trachypharynx (Vermidea) ... 552

Abdimia

sphenorhyncha, 509.

Aburria

carunculata, 523.

Acanthion

brachyura, 145.

Acanthodactylus

boskianus, 30.

Accentor

collaris, 471.

modularis, 471.

Accipiter

melanoleucus, 503.

nisus, 503.

pileatus, 503.

Aceros

nepalensis, 488.

Acheilostoma, gen. nov.,

549.

simpson, 549, 550.

Acomys

cahirensis, 5.

hunteri, 448.

Acridotheres

cristatellus, 481.

fuscus, 481.

gingimianus. 481.

mahrattensis, 481.

tristis, 481.

Acrocephalus

streperus, 471.

Acryllium

vulturinum, 528.

Addax

naso-maculatus, 1, 456. |

/Echmophorus

major, 932.

Abgialitis

curonica, 529.

hiaticula, 529.

7élureedus, 86.

A®lurus

fulgens, 442.

INDEX,

Aipyprymnus

rufescens, 464.

Aex

galericulata, 512.

‘sponsa, 512.

Agapornis

cana, 498.

pullaria, 498,

roseicollis, 498.

| Ageleeus

Srontalis, 481.

pheniceus, 481.

ruficapillus, 481.

Ajaja

rosea, 1, 510.

spinosa, 10.

Alactaga

decumana, 449.

indica, 449.

jaculus, 12.

Alauda

arborea, 483.

arvensis, 483.

celivor, 484.

crassirostris, 484.

cristata, 484.

gulqula, 484.

Alea

torda, 532.

_ Alcedo

| ispida, 64, 91, 488.

Alces

alces, 151.

machlis, 459.

Algiroides

africanus, 30.

nigropunctatus, 22.

Allactaga

longipes, 148.

Alma, 414.

| Alopex

lagopus, 135.

Proc. Zoou. Soc.+-1911, No. XXXVIT,

Amadina

bicolor, 475.

castanotis, 47.9.

erythrocephala, 4795.

Jasciata, 475.

Fringilloides, 475.

lathami, 475.

modesta, 475.

| Amblyrhamphus

holosericeus, 481.

Amblystoma, 104, 105,

115, 118.

tigrinum, 103.

Ammoperdix

honhami, 52).

heyi, 521.

Ampelis

cedrorum, 473.

garrulus, 473.

Amphipsylla, 388.

casis, 385, 386.

dea, 385, 386, 387.

Amyntas, 413.

Anas

boscas, 5138.

cristata, 513.

obscura, 513

pecilorhyncha, 513.

speculares, 514.

superciliosa, 514.

undulata, 514.

| Ancylostomum, 549.

| Anguilla, 183, 191, 192,

193, 194, 196, 216,

PAM 5 PS BY fe

vulgaris, 185, 186, 191,

192, 245.

Anisolabis

annulipes, 10.

Anoa

depressicornis, 454.

Anodorhynchus

glaucus, 495.

XXVI1 INDEX.

Anodorhynchus |) Aptenys: | Ardea

hyacinthinus, 495. mantelli, 51, d38: | gularis, 508.

leari, 495. owent, 93d. | herodias, 508.

Anourosorex Aquila | nove-hollandie, 508.

squamipes, 168. adalberti, AZ. | purpurea, 308.

Ans>r audax, 502. ralloides, 508.

albifrons, 512. | chrysaéius, 502. | sumatrana, 08.

brachyrhynchus, 512. | clanga, 502. | Ardetta

cinereus, 512. | imperialis, 502. | anvolucris, 508.

eyanoides, 512. | nevioides, 502. minuta, 5O8.

erythropus, 513. | werreauxt, 502. | Argoctenus, 297.

indicus, 513. | Avra aureus, 299, 306.

segetum, 513. ambiqua, 495. vittatus, 398.

A nseranas araraund, 86, 84, 85, | Argus

semipalmata, 512. 495. | giganteus, 522.

Anteliomys, 177. chloroptera, 495. | Artamus

Anthornis hahni, 495. | fuscus, 473.

melanura, 474. macao, 49. | personata, 473.

Antbracoceros maracana, 495. sordidus, 473.

coronatus, 488. militaris, 84, 495. | superctliosus, 473.

malabaricus, 488. | nobilis, 499. | Arvicanthis

malayanus, 488. severa, 495. pulchellus, 124, 148.

Anthropoides | Aramides ig s| striatus, 124, 148.

caruncilata, 527. ; cayennensis, 525. _ Arvicola

leucauchen, 83, 027. | ypecaha, 82, 525. | agrestis, 448.

leucogeranos, 527. | Aramus | amphibius, 147, 448.

paradisea, 4,53, 527. | —-scolopaceus, 528. | pratensis, 448.

virgo, 527. Araneus | terrestris, 147.

Anthropopithecus, 121, | verrucosus, 297, 299. | Asarcornis

358. | Arboricola | seutulatus, 4.

calvus, 429. | atrogularis, 520. | Asio

gorilla, 429. | torqueola, 520. | brachyotus, 492.

satyrus, 124, 125. | Arcella, 229. | mexicanus, 492.

troglodytes, 354, 429. Archzopsylla | ous, (45 72) 402:

Anthus erinacet, 365. _ Asopia

arboreus, 472. | sinensis, 865, 366. _ farinalis, 10.

obscurus, 472. | Archibuteo | Astur :

pratensis, 472. | dagopus, 501. | approximans, 68, 503.

spipoletta, 472. | Aretictis | nove-hollandie, 503.

Antilocapra | binturong, 181, 438. _ palumbarius, 68, 503.

americana, 458. | Arctogale | tachiro, 503.

Antilope | lewcotis, 438. | tibialis, 503.

cervicapra, 152, 456. | trivirgata, 438. | Asturina

Aphobus | Aretomys magnirostris, S02.

chopi, 481. caudatus, 446. natterert, 502.

Apodemus | himalayanus, 446. nitida, 502.

agrarius, 172, 173 | hodgsont, 416. | Ateles

— ningpsensis, 173. | marmotia, 446. | ater, 433.

— paliidior, 172, 173.

| monax, 446. | geoffroyi, 433.

fergussont, 172. | Ardea | grisescens, 433.

spectosus chevriert, 172. | agam, 508. | hybridus, 433.

sylvaticus, 148. | alba, 508. melanochir, 433.

Aprosmictus | argala, 49. paniscus, 433.

coccinopterus, 498. atricollis, 508. rufiventris, 433.

cyanopygius, 498. —— bubulcus, 508. variegatus, 433.

Aptenodytes | candidissima, 508. vellerosus, 433.

pennantt, 532. | einerea, 508. Athene

Apteryx, 55, 59, 88, 91, | coco, 508. brama, 493.

93. | egretta, 508. noctua, 72, 493.

australis, 51, 538. —— garzetta, 508. Atherina, 183, 204, 206,

haasti, 533. | goliath, 508. 209, 219.

Atherina

hepsetus, 205, 216, 247.

Atherura

africana, 449.

fasciculata, 439.

Atherurus

macroura, 14d.

Atlantoxerus

getulus, 149.

Aulacodus

swindernianus. 449,

Aulacorhamphus

sulcatus, Wo.

Babirussa

alfurus, 461.

babirussa, 140.

Balena

boops, 156.

muscilus, 156.

muysticetus, 155.

physalus, 156.

Balenoptera

boops, 156.

musculus, 156, 423.

physalus, 156.

Balearica, 82.

pavonina, O27.

requlorum, 527.

Balistes, 183, 215, 219.

capriscus, 209, 210,

248.

Bambusicola

Tytchii, 520.

thoracica, 520.

Barbastella

darjelingensis, 160.

Bassaricyon

alleni, 442.

Bassaris

astuta, 442.

Bassariscus, 137.

Baza

lophotes, 504.

Benhamia, 418.

austeni, 414.

Bernicla

brenta, 513.

canadensis, 513.

hutchinst, 518.

leucopsis, 513.

ruficollis, 513.

Bettongia

- cuniculus, 464.

gaimardi, 464.

lesueurt, 464.

penicillata, 464.

Bibos, 260, 264.

frontalis, 454.

INDEX,

_Bibos

gaurus, 54.

Bison

americans, 100, 454.

bison, Lot.

bonasus, Lod, 454.

Bitis

nasicornis, 2.

Biziura

lobata, 514.

Blarina, 166, 167.

Blarinella, gen. nov., 159,

166.

quadraticauda, 166,

167.

(Cryptotis) parva, 167.

Blastocerus

bezoarticus, 151.

Blennius, 204.

Bolborhynchus

lineolatus, 496.

Bos i

acutifrons, 256, 278,

211, 218, 209, 28).

bison, 154, 267.

bonasus, 154.

brachyceros, 249, 254,

256, 279, 280.

bubalus, 154, 251, 259.

chinensis, 264,

depressicornis, 250,

259!

Jrontosus, 249,

268, 282.

indicus, 154, 249, 278,

454.

264,

longifrons, 249, 254,

250, 262, Zid, 220;

281.

namadicus, 254, 256,

264, 272, 273, 277,

281.

planifrons, 26, 273.

primigenius, 249,

251, 254, 246, 258,

262, 264, 273, 277,

278, 279, 280, 281,

282.

taurus, 154, 249, 454.

— brachyceros, 254,

256.

— macroceros, 254.

— primigentus, 249,

256, 258, 281.

trochoceros, 250.

Boselaphus

tragocamelus, 456.

Botaurus

lentiginosus, 508.

stellaris, 508.

Bothrioneuron, 414.

200, |

Buceros

XXVI11

Box, 183, 219.

boops, 208, 216, 247.

Bradypus

didactylus, 132.

tridactylus, 132, 462.

| Breviceps, 397, 399.

sp., 405, 408, 411.

adspersus, 404.

gibbosus, 401, 404, 405,

406, 407, 408, 409,

410, 412.

macrops, 404.

mossambicus, 404.

panthert, 404,

verrucosus, 404.

Brotogerys

chrysopterus, 496.

jJugularis, 496.

pyrrhopterus, 496.

tirica, 496.

tut, 496.

tuipara, 496.

virescens, 496.

Bubalis

boselaphus, 454.

caama, 454,

tora, 454.

Bubalus, 260, 264.

equinoctialis, 454.

bubalis, 154.

buffelus, 4, 454.

caffer, 454.

Bubo

ascalaphus, 492.

bengalensis, 492,

blakistont, 492.

capensis, 72.

cinerascens, 71, '72.

coromandus, 492.

lacteus, 492.

maculosus, 71, 72, 492,

magel'anicus, 492.

maximus, 71, '72, 492.

nepatensis, 493,

orientalis, 498.

poensis, £93.

virginianus,

495.

lon tes

cylindrica, 60.

lunatus, 488.

rhinoceros, 488,

Bucorvus, 69.

abyssinicus, 61, 488.

cafer, 488.

Bungarus

Fasciatus, 4.

Buteo

albicaudatus, 50',

auguralis, 501,

berealis, 501.

al*®

XXV1IL

Buteo

desertorun, HUL.

erythronotus, DU1.

Ferox, SU.

jackal, 501.

pennsylvanicus, S01.

poliosomus, 5U\.

oulgaris, SUL.

Buteogallus

nigricollis, 502.

Butorides

atricapilles, 508.

cyanwrus, 508.

virescens, 508.

Bycanistes

subcylindricus, 483.

Cabassous

unicinctus, 41.

Cacatua

alba, 494.

citrina, 494.

ducorpsi, 494,

galerita, 495.

goffini, 499.

gymnops, 499.

hematuropygia, 495.

leadheateri, 495.

moluccensis, 495.

ophthalmica, 495.

roseicapilla, 499.

sanguinea, 499.

sulphurea, 84, 495.

triton, 495.

Caccabis

chukar, 521.

melanocephala, 52).

petrosa, 521.

rufa, 521.

saxatilis, 621.

Caica

leucogastra, 497.

melanocephala, 497.

axanthomera, 497.

Cairina

moschata, 512.

Calandrella

betiea, 484.

brachydactyla, 484.

Calearius

lapponicus, 480.

Calidris

arenaria, 529.

Callipepla

califormea, 521.

gambelli, 521.

squamata, 521.

Oalliste

brasiliensis, 474.

fastuosa, 474.

* Callorhinus

| Calopsittacus

INDEX.

Calliste

festiva, 474.

melanota, 474.

tricolor, 474.

Callithrix, 433.

jacchus, 127.

Callocephalon

galeatum, 84, 494.

ursinus, 133.

Calocitta

Jjormosa, 483. |

Calodromus, 57.

Caicenas

wicobarica, )18.

Calopelia

puella, 618.

nove-hollandia, 495.

Caluromys

philander, 143.

Calyptorhynchus

banksi, 494.

jfunereus, 494.

Cambridgea

antipodiana, 297, 300.

Camelus, 122.

bactrianus, 149, 150,

460. |

dromedarius, 149, 150, |

460. |

glama, 150. |

pacos, 150.

Cancroma

coehlearia, 508.

Canis

alopex, 134,

antarcticus, 440.

anthus, 440.

argentatus, 440.

aureus, 135, 440.

azare, 440. ~

bengalensis, 440.

eancrivorus, 440.

cerdo, 440.

chaina, 440.

dingo, 440.

Jamelicus, 440.

Familiaris, 134.

fulvicaudus, 440.

Sulvipes, 440.

Julous, 440.

griseus, 3.

hodophylax, 440.

hyena, 134.

gubatus, 440.

lagopus, 135, 440.

lateralis, 440.

latrans, 440.

leucopus, 440.

lupus, 134, 440. |

Canis

magellanicus, 440.

mesomelas, 440.

microtis, 440,

miger, 440.

niloticus, 440.

pallidus, 440.

pallipes, 440.

velox, 440.

virginianus, 440.

vulpes, 134, 440.

(Cuon) primevus,

440.

(—) rutilans, 440.

(Nyctereutes) pro-

cyonoides, 440.

Capra

egagrus, 457.

ammon, 153.

caucasica, 457.

cervicapra, 152.

depressa, 152.

dorcas, 153.

gazella, 152.

grimmia, 153.

hircus, 152, 457.

hispanica, 457.

ibex, 152, 457.

mambrica, 153.

megaceros, 457.

nubiana, 457.

pygmea, 152.

reversa, 152.

ruficapra, 152.

sinaitica, 457.

Capreolus

bedford, 180.

caprea, 459.

capreolus, 151,

pygargus, 459.

Caprimuleus

ewropeus, 487.

Capromys

brachyurus, 449,

pilorides, 449.

Caranx, 183, 219.

trachurus, 209.

Cardinalis

virginanus, 478.

Carduelis

elegans, 479.

orientalis, 479.

Cariacus

campestris, 151, 459.

gymnotis, 459.

leucurus, 459.

macrotis, 459.

mexicanus, 459.

nemorivagus, 459.

rufus, 459

virginianus, 489,

Cariama

cristata, 527.

Carphibis

spinicollis, 510.

Carpococcyx

radiatus, d4.

Carpodacus

erythrinus, 479.

hemorrhous, 479.

Carpophaga

e@nea 85, 516.

concinna, 8d.

latrans, 85.

paulina, 516.

Carvomys, 175.

Cassicus

hematorrhous, 480.

persicus, 480.

Cassidix

oryzivord, 60.

Castor

canadensis, 447.

jiber, 146, 447.

moschatus, 146.

Casuarius

australis, 51.

bennetti, 534.

bicarunculatus, 534.

casuarius, Dd4.

— australis, 534.

— beccarii, 534.

— intensus, d34.

— salvadori, 534.

— sclateri, 534.

— violicollis, 534.

intensus, 87.

papuanus, 534.

— edwardsi, 534.

philipt, 34.

picticollis, 534.

rothschildi, 87.

uniappendiculatus,

534.

— aurantiacus, 534.

— occipitalis, 534.

— rufotinctus, 524.

westermanni, O1, 87.

Cathartes, 61.

atratus, 50d.

aura, 69, 70, 505.

urubitinga, 50d.

Cavia

flavidens, 450.

porcellus, 146.

spixt, 450.

Cebus

albifrons, 432.

apella, 128, 432.

azare, 432.

capucinus, 128, 452.

fatuellus, 432.

INDEX.

Cebus

jlavescens, 432.

hypoteucus, 128, 432.

lunatus, 432.

monachus, 432.

vellerosus, 432.

versicolor, 432.

Centetes

ecaudatus, 444.

Centropus

phasianus, 490.

rufipennis, 490.

senegalensis, 490.

Centurus

tricolor, 487.

Cephalophus

abyssinicus, 2.

coronatus, 45.

dorsalis, 314, 455.

grimmé. 153, 409.

maxwelli, 455. -

monticolor, 455.

nigrifrons, 405.

rujilatus, +55.

sylvicultrix, 459.

Cepola, 183, 216,

218, 219.

rubescens, 212, 248.

Ceratodus, 118.

forsteri, 119.

Ceratogymna

elata, 488.

Ceratophyllus, 388.

crassus, 874, 379.

crispus, 365, 367, 370,

371.

dolabris, 371, 373, 374,

375.

euteles, 382.

Jtamulus, 380, 381.

fasciatus, 5, 9, 11.

lagomys, 871, 3872,

373.

londiniensis, 5.

mandarinus, 315, 377,

378.

mongolicus, 378.

pheopis, 383, 384.

tesquorwm, 379, 380.

Cerchneis

ardesiacus, 3.

Cercocebus

ethiops, 431.

albigena, 431.

aterrimus, 431.

collaris, 431.

chrysogaster, 431.

Juliginosus, 431.

hagenhecki, 431.

Cercoleptes

caudivolvulus, 442.

217

}

xkIK

Cercopithecus

ethiops, 128, 431.

albogularis, 430.

ascantas, 430,

barbatus, 126.

callitrichus, 431.

campbelli, 430.

cephus, 127, 430.

cynosurus, 431.

diana, 127, 480.

eryihrotis, 430.

grayt, 430.

griseoviridis, 123,

431.

lahiatus, 430.

lalandii, 431.

leucanepyx, 430.

Uhoesti, 430.

martin, 430.

mond, 430.

neglectus, 430.

mictitans, 430,

patas, 431,

petaurista, 430.

pygerythrus, 431.

pyrrhonotus, 431.

sabeus, 451.

schmidti, 430.

stairs?, 430.

talapoin, 431.

tantalus, 431.

Cereopsis, 513.

nove-hollandie, 512.

Ceriornis

blythi, 522.

caboti, 522.

melanoccphala, 522.

satyra, 522.

temmincki, 522.

Cerodon

rupestris, 450,

Cervicapra

bohor, 459.

isabellina, 455.

Cervulus

erinifrons, 458.

lacrymans, 458.

muntjac, 458.

reevesi, 458.

Cervus

atces, 151.

alfredi, 459.

aristotelis, 459.

axis, 459.

bezoarticus, 151.

camelopardatis, 150.

canadensis, 100,

458.

dama, 151.

davidianus, 459.

duvaucelli, 459.

XXX

Cervus

éelaphus,

458.

eldi, 459.

hippelaphus, 459.

kashmirianus, 458.

huhli, 459.

luehdorfi, 458.

mantchuricus, 459.

maral, 458.

moluccensis, 459.

philippinus. 459.

poreinus, 459.

schomburgki, 45%).

sika, 459.

taevanus, 459.

tarandus, 151.

Chalcopelia

chalcospilos, 518.

Chalcophaps

chrysochlora, 518.

indica, 518.

Chalcopsittacus

scintillatus, 494.

Chameepelia

passerina, 18.

talpacott, 518.

Charadrius

pluvialis, 529.

Charax, 208, 215, 219.

Chasmorhynchus

niveus, 484.

nudicollis, 484.

Chaulelasmus

streperus, 514.

Chauna

cristata, 511.

derbiana, 511.

Chen

cerulescens, 512.

hyperboreus, 512.

nivalis, 612.

Chenalopex

egypliaca, 513.

Jubatus, 2, 513.

Chenonetta

Jubata, 513.

Chera

progne, 476.

Chibia

hottentotta, 473.

Chinchilla

lanigera, 450.

Chionis

alba, 529.

minor, 529.

Chirogaleus

coquereli, 434.

milit, 434.

Chiromys

madagascariensis, 434.

150, 151,

INDEX.

Chlamydodera

maculata, 482.

Chloephaga

magellanica, 518.

melanoptera, 513.

poliocephala, 513.

rubidiceps, 513.

Chlorophonia

_spiza, 474.

viridis, 474.

Chiloropsis

aurifrons, 472.

hardwickii, 472.

Chodsigoa

berezowski, 166.

hypsibia, 166.

salenskii, 166.

smithii, 166.

Cholcepus

didactylus, 132.

Cholopus

didactylus, 462.

hoffmanni, 462.

Chrysochloris

asiatica, 142.

Chrysomitris

barbatus, 479.

spinoides, 479.

spinus, 479.

tristis, 479.

yarrellt, 479.

Chrysophrys, 215.

Chrysothrix

seiurea, 432.

Chrysotis

estiva, 496.

agilis, 496.

albifrons, 496.

amazonica, 496.

augusta, 496.

auripalliata, 496.

autwmnalis, 496.

bodini, 496.

bouqueti, 496.

brasiliensis, 496.

dufresniana, 496.

caymanensis, 496.

collaria, 496.

farinosa, 496.

Festiva, 496.

Jinschi, 496.

guatemale, 496.

guildingi, 496.

inornatus, 84.

leucocephala, 497.

levaillantt, 497.

mercenaria, 497.

ochrocephala, 497.

ochroptera, 497.

panamanensis, 497.

pretri., 497.

Chrysotis

rhodocorytha, 497.

salvini, 497.

ventralis, 497.

versicolor, 497.

vinacea, 497.

viridigena, 497.

vittata, 497.

wantholora, 497.

Chunga, 85.

burmeisteri,

O27.

Ciccaba, 72.

Nichlherminia

densirostris, 471.

Ciconia

alba, 509.

boyciana, 509.

nigra, 509.

Cineclus

aquaticus, 471.

Circaétus

gullicus, 503.

pecioralis, +.

Circus

eruginosus, AOL,

cineraceus, O01.

cyaneus, Oi.

gouldi, 501.

macrurus, 501.

maurus, 501.

Cissa

venatoria, 483.

Cissopis

leveriana, 475.

52, 73,

| Citellus

mongolicus, 379, 380.

Cittocincla

macrura, 471.

Clangula

glaucion, 514.

Clepsine, 419.

Cobboldia, 555.

Cobboldina, gen. nov.,

555.

vivipara, 555.

Cobus

ellipsiprymnus, 455.

anctuosus, 455.

Coccothraustes

melanurus, 478.

personatus, 478.

vulgaris, 478.

Cochoa

viridis, 473.

Celogenys

paca, 450.

Coendou

prehensilis, 145.

Coeereba

cyanea, 474.

Colaptes

auratus, 487.

Colivpasser

macrurus, 476.

Colius

capensis, 487.

castanotus, 487.

erythromelon, 487.

nigricollis, 487.

Culobus, 126.

vellerosus, 430.

Colugo

philippinensis, 130.

Columba

albiqularis, 516.

araucana, D16.

bolli?. 516.

caribbea, 516.

eversmanné, 16.

guined, 516,

gymnophthalma, 2, 4,

» Z

inornata, B16,

laurivora, 517.

leucocephala, 517.

leuconota, 517.

maculosa, D177,

ends, 217.

palumbus, 517.

picazuro, O17.

plumbea, 517.

rufina, 517.

speciosa, 517.

squamusa, O17.

Columbula

pieut, O17.

Colymbus

arcticus, 532.

glacialis, 532.

septentrionalis, I32.

Condylura

cristatus. 143.

Conepitus

mapurito, 441.

Connochzetes

gm, 454.

taurinus, 181.

Conuropsis

carolinensis, 496.

Conurus

acuticaudatus, 495.

eruginosus, 495.

aureus, 495.

auricapillus, 495.

aztec, 499.

cactorum, 495.

canicularis, 495.

chloropterus, 4995.

guarouba, 495.

hemorrhous, 495.»

INDEX.

Conurus

holochlorus, 495.

Jendayi, 495,

leucotis, 85.

nanday, 495.

ocularis, 495.

rubritorques, 496.

rubrolarvatus, 496.

solstitialis, 496.

wagleri, 496,

Copsychus

saularis, 471.

Coracias

affinis, 489.

garrulus, 489.

nevius, 4.

Coracopsis

barklyi, AX7.

nigra, 497.

vasa, 497,

Coreorax

melanorhamphus, 483.

Coregonus

acronis, 228.

Coris, 183, 217, 219.

giofred?, 208.

Qulis, 208, 209) 214,

216, 247.

vulyaris, 208.

Gorvina, 183, 215.

MWGhA 209 e210) eG:

217, 248.

Corvultur

albicoliis, 482.

Corvus

americanus, 482.

austratis, 482.

capellanus, 60, 482.

corar, 482.

cornix, 482.

corone, 482,

culminatus, 482.

daiiricus, 482.

Trugilegus, 482,

monedula, A82.

scapulatus, 482,

splendens, 482.

Coryexus, 282, 283.

africanus, 288, 294.

agilis, 290.

alatus, 287, 288, 289.

amazonicus, 288, 292,

293, 294.

andrewsi, 294, 295,

296.

anglicus, 289, 291,

294.

asiaticus, 288, 294.

catus, 287, 290, 296.

crassiusculus, 290.

XXxX1

Coryczeus

dane, 287, 290.

dubtus, 292, 296.

elongatus, 287, 238,

289.

erythreus,

293, 294.

jflaccus, 287, 288, 289.

288, 292,

Jurcifer, 287, 288.

gibbulus, 284, 285,

gracilicaudatus, 287,

290, 296.

hucleyi, 288.

latus, 287.

lautus, 287, 288, 294.

limbatus, 287, 289.

longistylis, 287, 288.

lubbocki, 288, 294.

megalops, 234, 285.

munrimus, 288, 294.

minutus, 287.

murrayt, 294, 296.

obtusus, 287, 289, 290,

291, 296.

ovalis, 287, 290, 291.

pacificus, 287.

pellucidus, 284, 285.

robustus, 287, 289.

speciosus, 282, 283,

287, 289.

tenuis, 288, 291, 292,

294, 296.

varius, 288.

venustus, 287, 289, 294.

vitreus, 287.

Corycelia, gen. nov.,

283.

sp., 287, 296.

brevis, 284, 235, 296.

carinata, 283, 284,

287,

concinna, 283, 284,

286, 287, 296.

curta, 284, 286, 296.

gibbula, 283, 284, 285,

286, 287.

gracilis, 283, 287.

longicaudis, 283, 284,

285.

pellucidus, 283, 284.

rostrata, 283, 284.

speciosus, 283.

tenuicauda, 283.

Coryphospingus

cristatus, 478.

pileatus, 478.

Coscoroba

candida, 512.

Cotinga

cincta 484

XXX1l

Coturnix

chinensis, 63, 520.

communis, 520.

coromandelica, 520.

histrionica, 520.

pectoralis, 520.

Cracticus

destructor, 483.

picatus, 488.

Cranorrbinus

corrugatus, 488.

Crax, 67.

alberti, 523.

alector, 5238.

carunculata, 55, 523.

daubentont, 523.

globicera, 56, 523.

globulosa, 528.

hecki, 56.

incommoda, 523.

sclateri, 523.

Crex

pratensis, 525.

Cricetomys

gambianus, 448.

Cricetulus

anderson, 173.

Cricetus

cricetus, 147.

frumentarius, 448.

Crithagra

albogularis, 479.

butyracea, 479.

chrysopyga, 479.

musicus, 479.

sulphurata, 479.

Crocidura

attenuata, 168.

cerulescens, 444.

Crocodilus, 87.

Crocopus

chlorogaster, 516.

phenicopterus, 516.

Crocuta, 93.

capensis, 96.

crocuta, 95.

gariepensis, 95,

germinans, 95, 99.

kibonotensis, 95, 98.

leonticwt, 95, 97, 98.

noltei, 95.

nyase, 99.

panganensis, 95, 92.

rufopicta, 97.

thierryt, 95.

thomasi, 98, 99.

togoensis, 95, 99.

wissmannt, 95, 96, 97.

Crossarchus

fasciatus, 1, 439.

obscurus, 439.

INDEX.

Crossoptilon

mantchuricum, 521.

tibetanum, 521.

Crotophaga

ant, 490.

Cryptobranchus, 117.

Cryptoprocta

ferox, 438.

Cryptotis, 167.

Crypturus, 93.

noctivagus, 5303.

obsoletus, 533

tataupa, 52, 58, 54,

533.

undulatus, 533.

Ctenocephalus

canis, 5.

felis, 5.

Ctenodactylus

gundi, 449.

Crenophthalmus, 388.

Ctenopsylla

mayardt, 9.

Cuculus

canorus, 490.

Curzeus

aterrimus, 481.

Cyanecula

suecica, 471.

Cyanocitta

coronata, 483.

cristata, 483.

melanocyaned, 483.

yucatanica, 483.

Cyanocorax

cyanomelas, 483.

cyanopogon, 483.

luxuosus, 488.

pileatus, 483.

Cyanolyseus

patagonus, 496.

Cyanopolius

cooki, 483.

cyanus, 483.

Cyanopsittacus

spixt, 495,

Cyanorhamphus

auriceps, 499.

matherbi, 499.

nove-zealandie, 499.

saissett, 499.

wnicolor, 499.

Cyanospiza

ciris, 477.

cyanea, 477.

Cyclopes

didactyla, 132.

Cygnus

atratus, 512.

bewicki, 512.

buccinator, 512.

Cygnus

musicus, 12.

nigricollis, 512.

olor, 512.

Cylicostomum, 551.

Cylindropharynx, gen.

noy., 5d1.

brevicauda, 551, 552.

longicauda, 551, 553.

Cynelurus, 135.

jubatus, 4, 438.

Cynictis

pemcillata, 439.

Cynocephalus

anubis, 432.

babouin, 432.

doguera, 432.

hamadryas, 482.

ibeanus, 432.

leucopheus, 432.

mormon, 432.

neumannt, 432.

papio, 432.

porcarius, 432.

sphine, 432.

volans, 124, 129.

Cynomys

ludovicianus, 446.

Cynonycteris

collaris, 444.

Cynopithecus

niger, 432.

Cynopterus

marginatus, 444.

Cyprinus

curpto, 228.

Cypselus, 60.

Cystophora

cristata, 448.

Dacelo

cervina, 488.

gigantea, 488.

leachi, 488.

Dacnis

cyana, 474.

Dafila

acuta, 514.

spuvicauda, 514.

Dama

mesopotamica, 459,

vulgaris, 459.

Damaliscus

albifrons, 454.

pygargus, 454.

Dasyprocta

acouchy, 450.

aguti, 450.

azare, 450.

Dasyprocta

cristata, 450.

isthmica, 450.

leporinus, 146.

mexicana, 450.

prymnolopha, 450.

punetata, 450.

Dasypus, 121, 122, 124.

minutus, 462.

novemcinctus, 141,

142.

quadricinetus, 141.

septemcinctus, 125,

142

sexcinctus, 141, 462.

tricinctus, 141.

unicinctus, 141.

villosus, 462.

Dasyurus

maculatus, 466.

viverrinus, 466.

Daulias

luscinia, 471.

Delphinus

delphis, 158.

orca, 158.

phocena, 158.

Dendraspis

viridis, 3.

Dendrocitta

himalayensis, 483.

sinensis, 483.

vagabunda, 423.

Dendrocopus

major, 487.

Dendrocygna

arborea, 513.

arcuata, 5138.

autumnalis, 513.

discolor, 54.

eytont, 513.

Sulva, 513.

javanica, 513.

major, 513.

viduata, 518.

Dendrolagus

bennetti, 464.

inustus, 464.

Deroptyus

accipitrinus, 497.

Desis

marina, 298, 299.

Desmana

moschatus, 146.

Dichoceros

bicornts, 488.

Dicotyles

labiatus, 461.

» tajacu, 461.

Didelphys

azare, 466.

INDEX,

Didelphys

cancrivorus, 466,

cinereus, 466.

crassicaudata, 466.

dorsigera, 144.

lanigera, 466.

marsupialis, 1438,

murina, 144, 466.

nudicaudata, 466.

opossum, 143, 466.

philander, 143, 466.

virginiana, 466.

Didunculus

strigirostris, 518.

Dignus

sowerbyi, 388.

Dilophus

carunculatus, 432.

Dioplodon, 422.

Diphyllodes

hunsteini, 2, 60.

_ Dipodillus

campestris, 449.

simont, 449.

Dipus

egyptius, 44%).

hirtipes, 449.

Dipylidium

caninum, 12.

Discoglossus

pictus, 408.

Dissura

episcopus, 509.

maguari, 50%).

Diuea

grisea, 477.

Dolichonyx

oryzivora, 480.

Dolichotis

patachonica, 450. —

| Dolomedes, 313.

tridentatus, 300, 311.

Donacola

castaneo-thorax, 476.

pectoralis, 470.

Dorcelaphus

americanus, 314.

— macrurus, 182.

Dorcopsis

ductuosus, 464,

miillert, 464.

| Drances

huttont, 313.

trippt, 313.

Drepanoplectes

jacksoni, 4.

Dromeus, 90.

nove-hollandie, 534.

Dromicia

nana, 464.

Xxx

Dryotriorchis

spectabilis, 508.

Dymecodon, 163.

Echidna

hystrix, 467.

Kelectus

cardinalis, 497.

pectoralis, 85, 498.

roratus, 498.

westermani, 498.

Ketopistes

migratorius, 517.

Himeria

(Coccidium) aviwm, 46.

Hlanus

ceruleus, 504.

: Elaphodus

michianus, 458.

| Hlephas

africanus, 452.

antiquus, 250.

indicus, 131, 42.

maximus, 124, 131.

Emberiza

aureola, 480.

ehrysophrys, 480.

cta, 480.

ciopsis, 480.

cirlus, 480.

citrinella, 480.

hortulana, 480.

luteola, 480.

melanocephala, 480.

miliaria, 480.

rutila, 480.

scheniclus, 480.

striolata, 480.

Entomyza

cyanotis, 474.

Kos

riciniata, 494.

reticulata, 494,

rubra, 494.

wallacet, 494.

Eothenomys, 175.

Epimys

andersont, 171.

confucianus, 170, 171.

— luticolor, 169.

edwards, 171.

excelsior, 170, 171.

jerdoni, 170.

ling, 170.

norvegicus, 170.

rattus, 147.

Equus

asinus, 155.

burchelli, 458.

caballus, 154.

grevyt, 458.

XXX1V

Equus

hemionus, 453.

-— indicus, 453.

hemippus, 453.

onager, 453.

quayga, 453.

somalicus, 453.

teniopus, 453.

zebra, 155, 4538.

Kremias

arguia, 24.

Hrethizon

dorsata, 145.

Erinaceus, 162.

albiventris, 444.

algirus, 444.

auritus, 444.

collaris, 444.

europeus, 142, 444.

frontatlis, 444.

micropus, 444.

miodon, 300.

Erithacus

rubecula, 471.

Erithizon

dorsatus, 449.

Erythrospiza

githaginea, 479.

Hrithrura

prasina, 476.

psittacea, 476.

Hsox, 189, 217, 219.

luctus, 190, 215, 243.

Estrelda

amadava, 479.

bella, 475.

bichenovit, 475.

cerulescens, 475.

cinerea, 479.

cyanogastra, 475.

dufresnit, 475.

Jormosa, 475.

granatina, 479.

melpoda, 475.

_phenicotis, 475.

phaéton, 475.

rubriventris, 475.

squamifrons, 475.

swbflava, 475.

temporalis, 475.

Eudocimus

albus, 510.

longirostris, 510.

ruber, 510.

Eudromias

morinellus, 529.

Eudynamis

orientalis, 65, 66, 490.

taitensis, 490.

Eudyptes

antipodum, 532.

INDEX,

Hudyptes

chrysocome, 532.

pachyrhynchus, 532.

selateri, 532.

Euphonia, 65.

chlorotica, 474.

jflavifrons, 474.

lanitrostris, 474.

nigricollis, 474.

pectoralis. 474.

sclateri, 474.

violacea, 60, 474.

Euphractus, 121, 124.

sexcinctus, 141.

Eaplectes

afer, 476.

capensis, 476.

jlammiceps, 476.

nigriventris, 476.

oryx, 476.

Euplocamus

albo-cristatus, 522.

andersont, 522.

erythrophthalmus, 522.

horsfieldi, 522.

lineatus, 522.

melanotis, 522.

nobilis, 522.

nycthemerus, O7, 8,

80, 522.

prelatus, 922.

swinhott, d22.

wretllott, 522.

Hupodotis

arabs, 526.

australis, 73, 74,

526.

caffra, 526.

denhami, 526.

kort, 4, 526.

rijicollis, 526.

Eupsychortyx

cristatus, 521.

leylandi2, 521.

sonnint, 57.

Hurypyga

helias, 528.

| Euspiza

americana, 480.

Exocetus

volitans, 228.

Faleo

esalon, 503.

biarmicus, 503.

concolor, 508.

eleonore, 503.

feldeggi, 69, 503.

jfusco-cerulescens, 503.

Falco

lanarius, 503.

melanogenys, 503.

peregrinus, 68, 503.

punicus, 503.

sacer, 503.

subbuteo, 503.

Felis

hengalensts, 436.

caffra, 437.

canadensis, 536.

caracal, 436.

catus, 136, 436.

chaus, 436.

chrysothri«, 436.

concolor, 436.

eyra, 437.

geoffroii, 437.

feo, 135, 314, 436.

lynx, 136, 436.

— rsabellinus, 181.

maniculata, 437.

nebulosa, 436.

onca, 136, 4386.

pardalis, 136, 437.

pardus, 135, 456.

passerum, 437.

planiceps, 436.

rubiginosa, 426.

rufa, 436.

serval, 436.

servalina, 436.

temminckt, 436.

tigrina, 437.

tiyris, 135, 436.

uncia, 456.

viverrina, 181, 436.

yaguarundt, 437.

Fiber

zibethicus, 448.

Fierasfer, 214, 219, 243.

acus, 210.

Foudia

erythrops, 477.

madagascariensis, 477.

Francolinus

bicalcaratus, 520.

cdpensis, 520.

clappertoni, 520.

coqui, 520.

granti, 520.

gularis, 520.

kirki, 520.

levaillanti, 520.

natalensis, 520.

pictus, 520.

ponticerianus, 520.

vulgares, 520.

Fratercula, 59. °

arctica, 78, 79, 80,

532.

Fregata, 89, 92.

aquila, 83, 506.

Fringilla

celebs, 6, 479.

kawarahiba, 479.}

linaria, 479.

montifringilla, 6, 479.

spodiogenia, 479.

teydea, 479.

tintillon, 479.

Fringillaria

sahara, 480.

Fulica

ardesiaca, 526.

atra, 526.

leucoptera, 526.

Fuligula

baert, 514.

cristata, 514.

JFerina, 514.

ferrugined, 514.

marila, 514.

rupina, 514.

Fulmarus

glacialis, 531.

Funisciurus

leucostigima, 2.

Furcifer

chilensis, 459.

Furnarius

rufus, 484.

Gadus,

219s 23.

morrhua, 211, 248.

Galago

allent, 484.

crassicaudata, 434.

garnetti, 434.

maholi, 434.

monteiri, 434.

Galeopithecus, 129.

philippinensis, 130.

Galeoscoptes

carolinensis, 471.

Galictis

barbara, 441.

vittata, 441.

Galidia

elegans, 439.

Gallierax

chlorochlamys, 491.

porphyreolopha, 491.

Gallinago

ce@lestis, 529.

Gallinula

chloropus, 526.

galeata, 526,

1835222; Zila;

INDEX,

Gallinula

nestotis, 526.

phenicura, 526.

pyrrhorhoa, 526.

| Galloperdix

lunulata, 520.

spadicea, 520.

zeylonensis, O=

Gallus

bankiva, 522

sonnerutt, 5

20.

stanteyt, 2.

varius, 52

Garrulax

albogularis, 472.

chinensis, 472.

leucolophus, 472.

pectoralis, 472.

perspicillatus, 472.

picticollis, 472.

Garrulus

glandarius, 483.

lanceolatus, 483.

Gasterosteus, 183,

PALS), Ai),

aculeatus, 204.

| spinachia, 201, 219.

| Gastropholis

vittata, 30.

Gazella

sp., 180.

arabica, 455.

bennett, 455.

cuviert, 45d.

dorcas, 153, 455.

euchore, 455.

loderi, 456.

marica, +56.

mohr, 456.

muscatensis, 456.

ruficollis, 1, 2.

rufifrons, 3, 456.

semmerringt, 456.

speki, 456.

subgutturosa, 456.

| Gecinus

viridis, 63, 64, 487.

Gelochelidon

anglica, 530.

Genetta

dongolana, 1.

Selina, 438.

genetta, 137.

pardina, 438.

senegalensis, 438.

tigrina, 438.

vulgaris, 438.

| Geocichia

citrina, 470.

wardi, 470.

202,

XXXV

Geococeyx

californianus, 95.

Geopelia

cuneata, 517.

humeralis, 517.

maugei, 517.

striata, 517.

tranquilla, 517.

. Geophaps

plumifera, 518.

seripta, 518.

Geopsittaeus

occidentalis, 499.

Georychus

capensis, 448,

Geotrygon

cristata, 518.

montana, 518.

mystacea, 518.

Geranoaétus

melanoleucus, 68,

502,

Gerbillus

egyptius, 448.

campestris, 448.

indicus, 448.

longifrons, 448.

pygargus, 448.

pyramidum, 448.

robustus, 448.

shawi, 448.

Giraffa, 458.

camelopardalis, 150.

Girardinus, 206.

Glareola

ocularis, 528.

pratincola, 529.

Glaucidium

passerinum, 493.

phalenoides, 493.

Globicera

aurore, d16.

pacifica, 516,

rufigula, 516.

Glossopsittacus

concinnus, 494,

Gobius, 183, 196, 204,

216, 218, 219, 235.

capito, 196, 203.

micropus, 41, 42, 48,

minutus, 40, 41, 42,43

44, 195, 196, 197,

219, 2389, 246.

niger, 195, 196, 197,

198, 200, 202, 203,

219, 246.

paganellus, 195, 196,

197,. 198, 210, 214,

Paste

XXXVI

Goura

coronata, 518.

victoria, 518.

Gracula

intermedia, 482.

Javanensis, 482.

religiosa, 60, 482.

Gr icupica

nigricollis, 482.

Grallina

australis, 472.

Grammatoptila

striata, 473.

Grayia

smyth, 181.

Grus

americana, 5217.

antigone, 5277.

australasiana, 527.

canadensis, 527.

communis, 527.

Japonensis, 527.

japonicus, 64, 65,

82.

monachus, 527.

virgo, 82.

Gubernatrix

cristatellus, 477.

Guira

piririgua, 490.

Guiraca

cerulea, 478.

cyanea, 478.

parellina, 478.

Gulo

gulo, 138.

tuscus, 139, 441.

Guttera

cristata, 522.

edouardi, 522.

pucherant, 523.

Gymnomystax

melanicterus, 4.

Gymnorhina

leuconota, 59, 60,

483.

tibicen, 413, 485.

Gymnura, 162.

Gypaétus

barbatus, 505.

Gypagus, 61.

papa, 69, 70, 505.

Gypohierax

angolensis, 505.

Gyps

bengalensis, 504.

Sulcus, 504.

kolbi, 504.

rueppelli, 504.

INDEX.

Heematopus

mger, 529.

ostralegus, 529.

Halcyon

chloris, 488.

sanctus, 64, 91, 488.

smyrnensis, 488.

vagans, 64, 488.

’ Haliaétus

albicillus, 502.

leucocephalus, 502.

leucogaster, 502.

leucoryphus, 502.

vocifer, 502.

Haliaster

tndus, 502.

intermedius, 502.

| Halicherus

gryphus, 443,

Hapale

chrysoleucus, 433.

Jacchus, 433.

melanura, 433.

penicillata, 433.

pygmed, 433.

Hapalemur

griseus, 434.

simus, 434.

Haplopelia

larvata, 518.

Harporbynchus

rufus, 471.

Harpyhaliaétus

coronatus, 66, S02.

Hedymeles

ludovicianus, 478.

Helictis

subaurantiaca, 441.

Helotarsus

ecaudatus, 5O2.

Hemicolea

rogenhoferi, 297, 298.

Hemidactylus

Jusciatus, 3.

Hemiderma

brevicauda, 130.

perspicillatus, 130.

Hemigalea

hardwickii, 438.

Hemitragus

jemlaicus, 457.

Hemixos

jlavala, 472.

Henicognathus

leptorhynchus, 496.

Herpestes

albicauda, 439.

auro-punctatus, 439.

Julvescens, 439.

galera, 439.

gracilis, 439.

Herpestes

griseus, 439.

ichneumon, 439.

pulverulentus, 439.

smithi, 439.

Herpetotheres

cachinnans, 504.

Neterhyphantes

melanogaster, 477.

Heterolocha

gouldi, 483.

Hieracicdea

berigora, 5O4.

Hierofalco

candicans, 503.

islandus, 504.

Himantopus

brasiliensis, 529.

nigricollis, 529.

Hippocampus, 208.

Hippopotamus

amphibius, 150,

461.

terrestris, 155.

Hippotragus

equinus, 456.

niger, 466.

Hirundo

rustica, 473.

Holaspis

guentheri, 30.

Homo

sapiens, 125.

troglodytes, 128.

Houbara

macqueent, 52, 73,

526.

undulata, 526.

Hyeena

brunnea, 439.

capensis, 9d.

crocuta, 93, Q4,

439.

cuviert, 95.

hyena, 134.

rufa, 90.

striata, 184, 439.

Wydrocherus

capybara, 490.

Hydromys

chrysogaster, 448.

Hydropotes

inermis, 409.

Hydrornis

alleni, 82, 525.

Hylobates

agilis, 353, 350, 356,

307, 429.

— martini, 399.

— piieata, 305.

gabrielle, 350.

Hylobates

hainanus, 353, 354,

355, 356, 357, 429.

hoolock, 355, 356, 357,

429.

lar, 355, 357, 429.

leuciscus, 353, 354, 355,

356, 357, 429.

leucogenys, 359, 429.

miilleri, 355, 429.

pileatus, 429.

syndactylus, 429.

Hylomys, 162.

Hymenolzemus

malacorhynchus, 514.

Hymenolepis

contracta, 15.

diminuta, 9, 10, 11.

horrida, 10.

mucrostoma, 13.

murina, 12, 13.

muris variegati, 13.

nana, 1d.

relicta, 10.

Hyomoschus

aquaticus, 460.

Hyphantornis

brachyptera, 477.

capensis, 477. ;

castaneo-fuscus, 477.

nigriceps, 477.

personata, 477.

superciliosus, 477.

textor, 477.

velatus, 477.

Hy pochera

mitens, 476.

Hypocolius

ampelinus, 473.

Hypophza

chalybea, 474.

Hypotriorchis

chicquera, 504.

Hypsipetes

maclellandi, 472.

Hyrax

capensis, 452.

dorsalis, 452.

Hystrichopsylla

tripectinata, 391.

Hystrix

brachyura, 145.

cristata, 144, 449.

dorsata, 145.

javanica, 449.

longicauda, 449.

macroura, 145.

prehensilis, 145.

Ibis

ethiopicus, 510.

INDEX.

Ibis

berniert, 510.

melanocephalus, \10.

strictipennis, 510.

Teterus

abeillei, 480.

baitimore, 480.

chrysocephalus, 480.

gamaici, 480.

spurius, 480.

tibialis, 480.

vulgaris, 480.

Icticyon

venaticus, 440.

Ictonyx

Jrenata, 441.

lybica, 441.

zorilla, 441.

Tonornis

martinicus, 525.

Isomys

variegatus, 448.

Ithaginis

geoffroyi, 520.

Ixocincela

crassirostris, 58, 54, 60,

Tynx

torquilla, 487.

Jaculus

jaculus, 148.

Julis, 215.

Junco

hyemalis, 480.

Ketupa

ceylonensis, 4933.

javanensis, 493.

Kobus

kob, 2.

unctuosus, 2.

Lacerta

31, 32, 40.

atlantica, 22, 26, 32.

boettgert, 25.

cerulea, 26.

campestris, 26.

chlorogaster, 22, 25.

defilippt, 25.

derjugini, 22.

dugesit, 22, 24, 25, 28, |

32.

echinata, 24,

32,

25, 30,

XxXxVvH

Lacerta

jfiumana, 26.

yalloti, 22, 25, 26;° 32.

horvathi, 34.

jacksoni, 19, 22, 23, 24,

30, 32, 34, 35, 36.

levis, 22, 24, 26, 32.

litfordi, 25, 31, 32.

mosorensis, 24, 25, 26,

27, 31, 34, 36.

muralis, 22, 23, 24, 26,

Ms} ill, Be, By, ais)

40,

— bocagit, 36.

— campestris, 25, 36.

— melissellensis, 26.

— monticola, 36.

— nigriventris, 22.

— serpa, 22, 25, 26,

— tiliguerta, 25.

— typiea, 23, 25, 51.

ocellata, 22, 24, 26, 28,

32.

oxycephala, 24, 25, 26,

27.

peloponnesiaca,

40.

pityusensis, 26.

sardoa, 39.

saxicola, 22, 2d.

schreibert, 347.

simony?, 22, 24, 25, 26,

32.

taurica, 37, 39, 40.

vauereselli, 33, 34, 35,

36.

viridis, 22, 24, 25, 26,

32, 37, 39, 40.

— gadovii, 347.

vivipara, 23, 24, 25, 31.

Lagopus

scoticus, 46, 100.

Lagostomus

trichodactylus, 450.

Lagothrix

humboldti, 433.

37,

| Lagotts

agilis, 22, 23, 24, 25, |

cuviert, 450.

Lama

glama, 150.

huanaeos, 460.

pacos, 150, 460.

peruana, 460.

Lamprocolius

auratus, 481.

chalybeus, 481.

Lampropsar

dives, 481.

Lamprotornis

eneus, 481.

XXXVI

Laniarius

quadricolor, 473.

Lanius

collurio, 473.

excubitor, 473.

lahtora, 473.

Larus

argentatus, 77, 030.

atricilla, 530. -*

brunneicephalus, 530.

cachinnans, 530.

caus, 531.

cirrhocephalus, 531.

dominicanus, 531.

Sranklini, 531.

Fuscus, 531.

gelustes, 531.

glaucus, 531.

wchthyaétus, O31.

leucopterus, 531.

marinus, 531.

nove-hollandia, 531.

ridibundus, 77, 531.

Latastia

degeni, 30.

hardeggeri, 30.

neumanne, 30.

phillipsii, 30.

spinalis, 30.

Latax

lutris, 138.

Leggada, 169.

Leipoa

ocellata, 523.

Lemmus

denuvmus, 146.

Lemur

albifrons, 434.

albimanus, 434.

brunneus, 434.

catta, 129, 434.

coronatus, 434.

flavifrons, 434.

Sulous nigrifrons, 5.

— rufifrons. 5.

macaco, 5, 434.

mayottensis, 434.

mongoz, 434.

niger, 434.

nigerrimus, 434.

wigrifrons, 434.

rujfitrons, 434.

rufipes, 434.

tardigradus, 129.

varius, 434. .

volans, 129, 130.

xanthomystax, 434.

Leontopithecus

amidas, 128.

edipus, 127.

Lepidosiren, i118, 119.

INDEX.

Leptobrachium, 393.

hasseltiz, 393, 395, +10,

403.

Leptoptila

jamaicensis, 018.

ochroptera, 518.

rufarilla, 518.

Leptoptilus

argala, 509.

crumeniferus, 509.

javanicus, 599.

Lepus

americanus, 451.

brasiliensis, 146,

451.

capensis, 145.

cuniculus, 145.

curopeus, 451.

nigricoilis, £01.

sechuenensis, 1&0.

sylvaticus, 451.

timidus, 145.

variabilis, 451.

Leucodioptron

canorum, 472.

Leuconerpes

candidus, 487.

Leucopternis

palliatus, 502.

Leucosarcia

picata, 518.

Licmetis

nasica, 495.

pastinator, 495.

Ligurinus

chloris, 478.

stnicus, 479.

Limnocorax

mger, 525.

Limnotragus, 350, 353.

Limosa

egocephala, 529.

lapponica, 529,

Linota

cannabina, 479.

flavirostris, 479.

rufescens, 479.

Liothrix

luteus, 471.

Lobivanellus

lobatus, 3, 530.

Lomvia

troile, 532.

Lophzemus

antareticus, 516.

Lophaétus

occipitalis, 503.

Lophophorus

impeyanus, 521.

thuyst, 521.

sclateri, 521.

Lophura

ignita, 314.

Loriculus

chrysonotus, 498.

galgulus, 498.

indicus, 498.

pusillus, 498.

Loris

gracilis, 434.

volans, 129.

Lorius

domicella, 494.

flavo-palliatus, 494.

garrulus, 494.

lory, 494.

tibialis, 494.

Lota

vulgaris, 229.

Loxia

bifasciata, 480.

curvirostris, 480.

leucoptera, 47).

pityopsittacus, 479.

Lucioperca, 215.

Lutra

Jelina, 181.

leptonyx, 441.

lutra, 138.

nair, 441.

vulgaris, 441.

Lycaon

pictus, 440.

Lycosa

canescens, 300.

Mabuia

raddonti, 181.

Macacus, 121, 125, 126.

arctoides, 431.

assamensis, 431.

brunneus, 4°41.

cyclopsis, 431.

cynomolgus, 431.

ecaudatus, 126.

inua, 126.

inuus, 126, 431.

lastotus, 481.

Teoninus, 431.

maurus, 431.

nemestrinus, 431.

ocreatus, 481.

pelops, 431.

philippinensis, 124,129.

pileatus, 431.

rhesus, 431.

rufescens, 431.

silenus, 126, 431.

stnicus, 431.

speciosus, 431.

syrichta, 129.

tcheliensis, 481.

oF ee.

Machetes

pugnax, 529.

Machetornis

rivosa, 484.

Macropodus, 207, 215.

Macropus

agilis, 464.

bennetti, AG4.

billardieri, 464.

brachiurus, 464.

derbianus, 464.

dorsalis, 464.

giganteus, 464.

gnelanops, 404,

parryi, 464.

robustus, 464.

ruficollis, 464.

rufus, 404.

thetidis, 464.

ualabatus, 464.

Macropygia

leptogrammica, 517.

phastanella, 517.

Manis

dalmani, 124, 1383.

pentadactyla, 124,

135.

tricuspis, 463.

(Pholidotis) gigantea, |

463. | Melursus

Manucodia

chalybea, 482.

Mareca

penelope, 514.

sibilatrix, 514.

Marmaronetta

angustirostris, J14.

Marmosa

dorsigera, 144.

murina, 144.

Marmota

marmota, 147.

monax, 147.

robusta, 169, 874, 375.

Martes

martes, 139.

zibellina, 139.

Megacephalon

maleo, 523.

Megaderma

spasma, 131.

Megalema

asiatica, 62, 490.

hodgsoni, 490.

virens, 490.

Megalophrys

(Leptobrachium) fee,

393-403.

montana, 393,

399.

nasuta, 393, 396, 399.

396,

INDEX.

Melanerpes, 60, 76, 89.

erythrocephalum, 487.

superciliaris, 62.

Melanocorypha

calandra, 48+.

mongolica, 454.

yeltoniensis, 484.

Meleagris

gallopavo, 522.

ocellata, 522.

Meles

ankuma, 441.

leptorhynchus, 44}.

meles, 140.

tarus, 441.

Melierax

monogrammicus, 503.

musicus, SUS.

polyzonus, 503.

Mellivora

capensis, 441.

indica, 441.

Melopelia

leucoptera, 517.

Melophus

melanicterus, 480.

Melopsittacus

undulatus,

499.

45, 8),

ursinus, 442.

Menura

superba, 484.

Mephitis

mephitica, 181, 441.

Mergus

albellus, 514.

merganser, d14.

serrator, 514.

| Meriones, 6.

Merops

apiaster, 489.

Mesoplodon

australis, 422.

bidens, 424.

densirostris, 421, 422.

haasti, 422.

(Dioplodon) — grayi,

420, 421, 422.

Mesopsylla

eucta, 12.

Meta

argentata, 298.

Metachirus

opossum, 1438.

Metopiana

peposaca, 51+.

Metriopelia

melanoptera, 518.

Microcebus

smithi, 434.

XXX1X

| Microglossus

aterrimus, 494,

Microsittace

Serruginea, 496.

Mierotus

arvalis, 178, 175.

calamorum, 174, 175.

inez, 176.

irene, 173.

limnophilus, \74, 175.

malcotmt, 178, 174, 175,

178.

nuc, 176.

ratticeps, 175.

saved, 170.

(Anteliomys) chinensis,

175.

(Caryomys) eva, 175.

(Eothenomys) melano-

gaster, 176.

Midas

chiysomelas, 433.

geoffrott, 433.

labiatus, 433.

mystax, 433.

edinus, 433.

rosalia, 433.

rujimanus, 433.

ursulus, 433.

Milvago

australis, 504.

chimachima, 4, 504.

chimango, 504. .

megalopterus, 504.

Milvulus

tyrannus, 484.

Milvyus

egyptius, 504.

govinda, 504.

actinus, 504.

migrans, S04.

Minus

polyglottus, 471.

saturinus, 471.

Mirafra

affinis, 484.

cantillans, 484.

Miro

albifrons, 4771.

Mirounga

leonina, 133.

Mitua

tomentosa, 523.

tuberosa, 523.

Molothrus

badius, 481.

bonariensis, 481.

purpurascens, 48}.

Momotus

brasiliensis, 489.

subrufescens, 489.

Monachus

albiventer, 445.

Monodon

monoceros, 19d.

Monticola

cyanus, 470.

saxatilis, 470.

Montifringilla

nivalis, 479.

Morphnus

guianensis, 502.

Moschus

moschiferus, 150,

458.

sifanicus, 180.

Motacilla

flava, 472.

lugqubris, 472.

melanope, 472.

rail, 472.

Motella, 215.

mediterranea, 207.

Mungos

ichneumon, 136.

Munia

castanetthorax, 101.

ferruginea, 475.

flaviprymna, 101.

maja, 476.

malaharica, 476.

malacca, 476.

nisoria, 476.

punctularia, 476.

rubro-nigra, 476.

striata, 476.

topela, 476.

Murena, 216, 219.

Mus

amphibius, 147.

arboricola, 448.

evellanarius, 147.

éandicota, 448.

barbarus, 448.

blanfordi, 448.

cervicolor, 448.

cricetus, 147.

exulans, 448.

gansuensis, 169.

gaculus, 148.

lemmus, 146.

leporinus, 146.

longipes, 148.

marmota, 147.

sninutus, 448.

monax, 147.

musculus,

147.

porcellus, 146.

ratius, 4, 147, 448.

striatus, 148.

sylvaticus, 148, 448.

122, 146, |

_ Mynoglenes

INDEX.

Mus

terrestris, 147.

volans, 148, 149.

Museardinus

avellanarius, 147, 447.

Muscicapa

atricapilla, 473.

Musophaga

TOSS@, 3.

violacea, 491.

Mustela

barbara, 188.

bed fordi, 169.

dorsalis, 169.

erminea, 138, 139,

441,

flavigula, 181.

foina, 441.

Furo, 139.

gulo, 138.

kathiah, 168, 169.

lutra, 138.

lutris, 138.

martes, 139, 441.

nivalis, 168.

pennanti, 441.

putorius, 139, 441.

rixosa, 168, 169.

russellianda, 168.

vulgaris, 138, 441.

zibellina, 139.

Mustelis

diversis, 122.

Mycerobas

melanoxanthus, 478.

Mycetes, 453.

Mycteria

americana, 509.

Myiophones

horsfieldt, 470.

chiltont, 299,

309.

insolens, 297.

marrinert, 310.

Myodes

lemmus, 448.

Myopotamus

coypu, 449.

Myopsittacus

monachus, 496.

Myospalax

cansus, 178, 179.

—— shenseius, 178.

fontaniert, 391.

Myotalpa

vufescens, 179.

Myotis

altarium, 161.

bechsteini, 161.

moupinensis, 162.

308, |

Myotis

pequinius, 161.

Myoxus

dryas, 447.

gtis, 447.

quercinus, 447.

_ Myristicivora

luctuosa, 516,

Myrmecophaga

didactyla, 132.

jubata, 124, 132, 462.

tetraductyla, 133.

tridactyla, 124, 132.

Myrus, 113, 189, 192,

193, 216, 217, 218.

219.

vulgaris, 191, 192.

Myzantha

garrula, 474.

Naia

tripudians, 4.

Nandinia

binotata, 439.

Nanodes .

discolor, 499.

Nasua, 137.

naréica, 442.

rufa, 442.

| Neeator, 549.

Necturus, 103, 105, 115,

119.

Nemorhedus

goral, 456.

' Neopbema

bourkti, 499.

chrysogastra, 499.

elegans, 499.

pubchella, 499.

splendida, 499.

venusta, 499.

Neophron

percnopterus, 504.

| Neopsylla

aliena, 388, 390.

bidentatijormis, 387,

388.

compar, 387.

Neotetracus, 159.

sinensis, 162.

Neotragus

pygmea, 152.

Nephila

argentatum, 298.

venosa, 299.

Nerophis, 183, 201, 219,

225, 235.

equorius, 199, 200,

248.

dn Sght

a ee

were

Nesochen

sand vicensis, 515.

Nesonetta

aucklandica, 514.

Nesopelia

galapagoensis, 517.

Nestor

notabilis, 84, 494.

Ninox

boohook, 71, '72, 493.

nove-scelandia, 493.

Nipponia

temmincki, 510.

Nisaétus

Jasciatus, 502.

pennaéus, S02.

spilogaster, 502.

Noctilio

leporinus, 131.

Nothoerax

urumutum, 523.

Nothura

. maciulosa, 535.

Notiosorex, 167.

Nucifraga

caryocatactes, 482.

Nucras

delalandii, 30.

Numenius

arquata, 529.

Jemoralis, 529.

pheopus, 529.

Numida

ellioti, 522.

meleagris, 522.

mitrata, 522.

ptilorhyncha, 522.

reichenowi, 522.

Nyctea

scandiaca, 4938.

Nycticebus

Javanicus, 434.

tardigradus, 434.

Nycticorax

caledonicus, 508.

gardeni, 508.

griseus, 508.

vielaceus, 4, 508.

Nyctidromus, 60.

Nyctipithecus

Jfelinus, 433.

rufipes, 433.

trivirgatus, 433.

vociferans, 435.

Nyinphicus

cornutus, 499.

uveensis, 499.

Ochotona

cansa, 180.

INDEX,

Ochotona

hodysoni, 180.

tihelana, 180.

Octodon

cumingi, 449.

Ocydromus

australis, 525.

earli, 25.

Suseus, 5295.

lafresnayanus, 25.

sylvestris, 525.

Ocyphaps

lophotes, 518.

Odobenus

rosmarus, 134,

Odontophorus

dentatus, 521.

guianensis, 521.

Gidemia

nigra, O14.

Gidicnemus

bistriatus, 529.

grallarius, 77, 529.

scolopax, 76, 77, 529.

superciliaris, 529.

Gina

capensis, 518.

Onychogale

Jrenata, 464.

unguifera, 464.

Ophichthys, 183,

219.

(Sphegebranchus) 77-

189, |

berbis, 192, 193, 216,

217, 245.

(Ophisurus) — serpens,

193, 216.

Ophidium, 183, 205, |

206, 214, 215, 216, |

Qe 2NGy 220!

barbatum, 197,

210, 247.

Orchesticus

ater, 475.

Orcinus

orca, 158.

Oreas

canna, 456.

Oreotragus

saltator, 455.

Oriolus

indicus, 473.

kundoo, 473.

melanocephalus, £73.

Ornithorhynchis

paradoxus, 181.

Ortalis, 58.

alhiventris, 528.

garrula, 523.

motmot, 523.

rujicauda, 56, 523.

Proc. Zoou. Soc.—1911, No. XX XVIII.

204, |

Ortalis

vetula, 523,

Ortyx

virginianus, 521.

Orveteropus

ethiopicus, 465.

capensis, 403.

Oryctolagus

cuniculus, 145.

Oryx

algazel, 1.

beatriz, 456.

beisa, 456.

gazella, a2.

leucoryx, 456.

Oryzoborus

crassirestris, 478.

torridus, 478.

Osmotreron

bicineta, 516.

vernans, O16.

Ostinops

decumanus, 480.

Otaria

culiforniana, 443.

hookeri, 443.

Jubata, 445.

pusilla, 443.

Otis 85.

tarda, 73, 526.

Otocorys

alpestris, 43+.

hilophu, 484.

Otocyon

megalotis, 440.

Ouaearia, 435.

Ourebia

montana, 1.

nigricaudata, 2, £55.

Ovis

ammon, 133.

aries, 195.

blanfordi, 457.

burrhel, 457.

cycloceros, 457.

guineensis, 153.

hodgsoni, 457.

musimon, Ad7.

strepsiceros, 153.

tragelaphus, 457.

vignei, 407.

Pachynus

bruchyurus, 2, 497.

Padda

oryzivora, 476.

Pagellus, 215.

Pagophila

eburned, O31.

Palzopsylla, 388.

Palzornis

alexandri, 498.

caniceps, 498.

cyanocephala, 498.

docilis, 498.

fasciata, 498.

longicauda, 498.

nicoburica, 498.

peristercdes, 498.

rosa, 498.

torquatus, 498.

Palamedea, 86.

cornuta, 511.

Pandion

haliaétus, 501.

Panurus

biarmicus, 471.

Papio

cynamolgos, 128.

hamadryas, 127,

128.

Paradisea

minor, 482.

raggiana, 60.

rubra, 2, 482.

Paradisornis

rudolphi, 60.

Paradoxurus

aureus, 438.

griayt, 438.

hermaphroditus, 438.

larvatus, 438.

leucomystax, 438.

niger, 438.

pallast, 438.

philippensis, 438.

prehensilis, 438.

Pardosa, 298.

canescens, 300.

Pareiasaurus, 118.

Pareudrilus, 415. i

pallidus, 419.

Paroaria

capitata, 477.

cucullata, 477, 485.

larvata, 477.

Parus

ater, 471.

borealis, 471.

ceruleus, 471.

caudatus, 471.

cristatus, 471.

eyanus, 471.

major, 471.

palustris, 471.

varius, 471.

Passer

alario, 48.

arcuatus, 478.

diffusus, 478.

INDEX.

Passer

luteus, 478.

montanus, 478.

salicicola, 478.

simplex, 478.

Pastor

roseus, 482.

Pauxis

galeata, 523.

Pavo

cristatus, 522.

nigripennis, 522.

spicifer, 522.

Pelecanus, 83.

conspicillatus, 506.

crispus, d06.

Suscus, 506.

manillensis, 506.

mitratus, 506.

onocrotalus, 506.

rufescens, 506.

trachyrhynchus, 3.

| Pelobates, 394.

Pelodytes

punctatus, 397.

Penelope

cristata, 523.

greeyt, 923.

Jacucaca, 623.

mara, 523.

pileata, 523.

purpurascens, 523.

superciliaris, 58, 523.

Penthetria

laticauda, 4.

Peragale

lagotis, 466.

Perameles

obesula, 466.

Perea, 183, 210,

215,

216, 219, 227, 237, |

240.

fluviatilis,

248.

Perdicula

argqoondah, 521.

asiatica, 521,

Perdix

cinerea, 520.

hodgsome, 520.

Perichxta, 412.

Peristedion

cataphractum, 201, 202,

210, 246.

Peristera

geoffroyt, 518.

Peristethus, 183,

201, 208, 216, 219.

214, 229

195,

cataphractus, 201, 202,

210, 219, 246.

Pheretima,

Pernis

apivorus, 501.

ptilorhyncha, 501.

Perodicticus

potto, 434.

Petaurus

australis, 464.

breviceps, 464.

sciureus, 464.

Petrogale

brachyotis, 464.

penicillata, 464.

xanthopus, 464.

Petronia

petronelia, 478.

stulta, 478.

Phacochcerus

ethiopicus, 461.

africanus, 461.

Phaéthusa

magnirostris, 530.

Phalacrocorax, 83.

africanus, 507.

brasilianus, 507.

carbo, 507.

carunculatus, 507.

graculus, 507.

javanicus, 507.

lugubris, 507.

pygmeus, 3.

varius, 507.

Phaps

chalcoptera, 518.

elegans, 518.

histrionica, 518.

Phascolarctos

cinereus, 464.

Phaseolomys

latifrous, 465.

mitchelli, 405.

Phasianus

colchicus, 52).

decollatus, 521.

ellioti, 621.

principalis, 521.

reevesi, 521.

semmerring?, O21.

torguatus, 521.

versicolor, 521.

wallichiti, 521.

412, 418,

419.

cingulata, 413.

montana, 418, 414, 416,

417.

Pheucticus

chrysogaster, 478.

Phlogeenas

luzonica, 918.

slairt, 18.

INDEX, x]

Phoca Platycercus | Poephagus

leonina, 133. adelaide, 498. | grunniens, 454,

rosmarus, 134. barnardi, 499, Poéphila

brownt, 499. |

elegans, 499.

acuticauda, 476.

cincta, 476.

ursina, 133.

vitulina, 1383, 134,

443, erythropeplus, 499. | gouldie, 476.

Phocrna eximius, 499. mirabilis, 476.

phocena, 158. | jflaveolus, 84, 499. Polioaétus

Phodilus | flaviventris, 499. plumbeus, 502.

badius, 492. | acterotis, 499. Poliopsar

Pheenicopterus pallidiceps, 499. andamanensis, 481.

pileatus, 499.

semitorquatus, 499. matlabaricus, 481.

zonarius, 499. Polyacanthus, 207,

Pleeotus | ZAlay

ignipalliatus, 511. blythi, 481.

roseus, O11,

rubra, O11.

Phonipara

canora, 477. egyptiacus, 160. Polyboroides

olivacea, 477. ariel, 160. typicus, 504.

Phoronis, 233. auritus, 131, 160, 4465. Polyborus

Phrygilus, 477. christiei, 160. brasiliensis, 504.

alaudinus, 478. homochrous, 160. Polyplectron

Sruticeti, 478.

gay, A478.

bicalcaratum, 522.

chinguis, 522.

sacrimontis, 160. |

wardi, 160, 161.

Physeter Plectrophenax germaint, 522.

catodon, 157. nivalis, 480. Polytelis

macrocephalus, 157. Plectropterus alexandre, 498.

barrabaudi, 498.

melanurus, 498.

gambensis, 512.

niger, 512.

microcephalus, 157.

microps, 17.

tursio, 157. rucppelli, 512. Porcula

Pica Plegadis | salvania, 461.

bootanensis, 482. falcinellus, 510. | Poromera

mauritanica, 482, guarauna, 510. | fordiz, 30.

rustica, 482. Ploceus Porphyrio

sericea, 482. atrigula, 477. | albus, 82.

Pinicola bengulensis, 477. ceruleus, 525.

enucleator, 479. manyar, 477. calvus, 525.

Pionopsittacus Plotus edwardsi, 525.

pileatus, 497. anhinga, 3, 507. madagascariensis, 82,

Pionus levaillant?, 507. 52.

melanonotus, 82, 525.

poliocephalus, 525.

vitiensis, 525.

Porrhothele

chalcopterus, 497. melanogaster, 507.

Suscus, 497. Pluvianus, 73, 76, 77, |

maximiliant, 497. 89.

menstruus, 497. egyptius, 74, 75. |

senilis, 497. | Podager | antipodiana, 298.

sordidus, 497. nacunda, 314. Porzana

Pipile Podargus carolina, 525,

cujubi, 523. cuviert, 58, 60, 489. | maruetta, 525,

cumanensis, 528, Podiceps | notata, O29.

jacutinga. 523. cristatus, 78, 81, Potamocherus

Pipilo 532. africanus, 461.

erythrophthalmus, 480. | _ griseigena, 532. penicillatus, 461.

Pitangus | Pecilonetta poreus, 140.

Potorous

tridactylus, 464.

Pratincola

rubetra, 471.

sulphuratus, 484.

Pithecia, 433.

Pithecus, 121, 125.

Pitta

bahamensis, 514.

erythrorhyncha, 514.

Peocephalus

fuscicapillus, 497.

bengalensis, 484.

strepitans, 484.

Pitymys, 173.

Platalea

alba, 510.

leueorodia, 510.

‘fuscicollis, 497. | Presbytis, 124.

gulielmt, 497.

meyert, 497.

robustus, 497.

rueppelli, 497.

senegalus, 497.

aygula, 124, 127.

mitratus, 124, 127.

Procyon

cancrivorus, 442.

lotor, 140, 442.

xliv

Proedromys,

WAT

hedtordi, 159, 177.

Prosthemadera

nove-zealandie, 474.

Procteles

cristatus, 439.

Protopterns, 118, 119.

Psammomys

obesus, 448.

159, 174,

Psephotus

chrysoprerygius, 499.

hematonotus, 84,

499.

multicolor, 499.

pulcherrimus, 499,

Pseudis, 396.

Pseudochirus

peregrinus, 464.

- Pseudoleistes

virescens, 481.

Pseudotantalus

this, 5U9.

leucocephalus, 509.

Psittacula

passerina, 496.

Psittacus

erithacus, 497.

timneh, A97.

Paittenteles

chlorolepidotus, 494.

Psittinus

incertus, 498.

Psophia

crepitans, 27.

leucoptera, 527.

viridis, 527.

Pternistes

afer, 520.

anfuscatus, 520.

leucoscepus, 520.

nudicollis, 420.

swainsoni, 520.

Pterocles

alchata, 519.

arenarius, 519.

bicinctus, 519.

exustus, 519.

Jasciatus, 519.

namaqua, 519.

seneqallus, D9.

Pteroglossus

inscriptus, 490.

torquatus, 490,

wiedi. 490.

Pteromys

albo-rufus, 446.

enornatus, 446.

lencogenys, 446.

magnificus, 446,

oral, 446.

INDEX.

Pteropus

edulis, 130.

JSormosus, 444.

gouldi, +44.

medius, 444.

poltocephalus, 444.

pstlaphon, 444.

scapulatus, 444.

vampyrus, 130.

Plonorhynehus

smithi, 482.

wiolaceus, 60, 482.

Ptilopachys

ventralis, 520.

Ptilopus

bellus, 85.

Susciatus. 516.

jambu, 516.

melanocephalus 516.

porphyraceus, 516.

Ptilorhis

paradisea, 482.

Ptilostomus

senegalensis, 483.

tistes

erythropterus, 493.

_ Pucrasia

darwini, 521.

macrolopha, 321.

axanthospila, 521.

Pudua

humilis, 459.

Puffinus

anglorum, 531.

kuhla, 531.

major, O31.

Pulex

irritans, ).

Pulsatrix

torquata, $93.

Pycnonotus

atricapillus, 472.

crocorrhous, 472.

hemorrhous, 472.

Jocosus, 472.

leucotis, 472.

sinensis, 472.

xanthopygos, 472.

Pygathbrix, 124, 127, 128.

Pygosceles

tematius, 532.

Pyranga

sairad, 475.

Pyrenestes

albifrons, 477.

Pyromelana _

aurea, 476.

JSranciscana, 477.

Pyrrhocorax

alpinus, 483.

graculus, 488.

Pyrrbula

europea, 479.

rubicilla, 479.

Pyrrhulauda

verticalis, 484.

Pyrrbulopsis .

" personata, 498.

splendens, 498.

tabuensis, 498.

Pyrrhura

cruentaia, 496.

leucotis, 496.

perlata, 496.

vittata, 496.

Physeter

catodon, 124.

macrocephulus, 124.

Pytelia

citerior, 475.

phenicoptera, 475.

wienert, 475.

Quelea

occidentalis, 477.

russt, 477.

sanguinirostris, 477.

Querquedula

brasiliensis, 514.

castanea, 514.

circia, 514,

erecea, 5A.

eyanoptera, 514.

Faleata, 514.

flavirostris, 514.

formosa, d14.

gibberifrons, 514.

Quiscalus

lugubris, 481.

versicolor, 481.

Rallina

pectloptera, 525.

Rallus

abboti, 82.

aquaticus, 525.

celebensis, 525.

maculatus, 925,

pectoralis, 5265.

rhytirhynchus, 525,

Rana, 401.

Rangifer

tarandus, 151, 459.

— caribou, 181, 182.

Raphicerus

melanotis, 455.

tragulus, 459.

Recurvirostra, 69.

avovetta. 77, 78, 429.

Rbhamphastos

ariel, 68, 489.

brevicarinatus, 489.

carinatus, 63, 489.

cuviert, 489.

discolorus, 489.

erythrorhynchus, 490.

tocard, 490.

toco, 490.

vitellinus, 490,

Rhampboceelus

brasilius, 475.

Rhamphocorys

clotheyi, 484.

Rhea, 90, 93.

americana, 51, 52,

04, 535.

darwin, 535.

macrorhyncha, 535.

Rhinoceros

bicornis,

453.

lasiotis, 453.

sondaicus, 453.

sumatrensis, 453.

unicornis, 144,

453.

Rhinochetus

Jubatus, 528.

Rhinoderma

darwini, 401.

Rhinolophus

cornutus

160.

Rhizomys

badius, 449.

davidi, 179, 180.

sinensis, 179.

vestitus, 179.

Rhodonessa

caryophyllacea, 512.

Rhynchotus

perdicarius, 533,

rufescens, 533.

Rhytidoceres

plicatus, 488.

undulatus, 488.

144,

pumilus,

Rissa

tridactyla, 531.

Rollulus

eristatus, O21.

Roatrbamus

sociabilis, 504.

Rupicapra

rupicapra, 152.

tragus, 456.

Rupicola

crocea, 2, +84.

Ruticilla

phenicurus, 471.

tithys, 471,

INDEX,

| Saimiri

morta, 128.

seiurea, 129.

Salamandra

atra, 347.

maculosa, 823, 324,

O20) 820), S28; 329,

BB0) Sell, vor, ood)

340, 342, 347.

— algira, 324, 330.

— coccinea, 338.

— corsica, 324,

340, 342, 343.

— europea, 324.

— gallaica, 327, 335, |

334, 335, 335.

— molleri, 324, 327,

dan, Go4, Gan

337, 342, 347.

— nigriventris, 324.

— quadrivirgata, 324.

—teniata, 324, 327,

Sal), why,

338, 339,

347.

Saltator

aurantitrostris, 475,

magnus, 479,

similis, 475.

Sarcidiornis

carunculata, 512.

melanonota, 3,

512.

Sarciophorus, 89.

_pectoralis, 76, 529.

Sarcophilus

satanicus, 100.

ursinus, 466.

Sarcorhamphus

equatorialis, 505,

gryphus, 2, 506.

Sargus, 183, 208, 215,

219.

rondeletii, 209, 210,

211, 216, 248.

Sauropatis, 64.

Saxicola

enanthe, 471.

Sealops

aquaticus, 143,

Scapteira

knowit, 31.

Seardafella

sguamosa, 517,

Scaurus

striatus, 10.

Sceloglanx

albifacies, 493.

Scizena, 215.

Sciuropterus

jimbriatus, 446,

Spie}, Geiig |

345, 346,

xIv

Sciuropterus

layardi, 446.

russicus, 149

volans, 148, 446.

volucella, 446.

Scinrotamias

davidianus, 169.

— consobrinus, 169,

3871, 3884, 380,

393,

Sciuras

éstivans, 445.

arizonensis, 445,

atrodorsalis, 45.

bicolor, 445.

caniceps, 445.

castaneiventris, 445.

cinereus, 149, 445.

dorsalis, 445.

ephippium, 445.

erythropus, 445.

flavus, 149.

getulus, 149.

griseo-flavus, 445,

griseus, 445.

hudsonicus, 445.

hypopyrrhus, 445,

indicus, 445.

ludovicianus, 445.

macrurus, 445,

madagascariensis, 445.

maximus, 445.

niger, 149.

nigro-vittatus, 445,

palmarum, 445.

plantani, 445.

prevosti, 445,

punctatus, 2.

pusillus, 445.

pygerythrus, 445.

stramineus, 440.

striatus, 149.

syriacus, 445,

tennenti, 445.

tristriatus, 445,

variabilis, 446.

variegatus, 446.

volans, 149.

vulgaris, 148, 446.

vulpina, 446.

Scolopax

rusticola, 529.

Scops

asio, 493.

git, 493.

glabripes, 493.

lempiji, 493.

leucotis, 493.

senegalensis, 493.

Scopus

umbretia, 3, OO.

xlvi

Scotopelia

bouviert. 71.

peli, 493.

Scythrops

nove-hollandie, 66.

Selenidera

maculirostris, 490.

Seleucides

nigricans, 482.

Semnopithecus

cephalopterus, 430.

entellus, 430.

fasciatus, 430.

hypoleucus, 450.

maurus, 430.

obscurus, +30.

Sericulus

melinus, 470.

Serinus

angolensis, 479.

canarius, 479.

canicollis, 479.

hortulanus, 479.

totius, 479.

Serpentarius

reptilivorus, OD.

Serranus, 183, 215, 219.

cabrilla, 209.

Sialia

wilsont, 471.

Sibia

capistrata, 472.

Simia

e@thiops, 128.

apedia, 126.

apella, 128.

aygula, 127.

capucina, 128.

cephus, 127.

cynamolgus, 128.

diana, 127.

faunus, 126.

hamadryas, 127.

Jacchus, 127.

midas, 128.

morta, 128.

nemed, 127.

edipus, 127.

paniscus, 127.

satyrus, 121, 122, 124,

125, 429.

sciurea, 129.

sidenus. 126.

sphine, 126.

sylvana, 121, 122, 124,

125, 126.

syrichta, 129.

Siphonostoma, 183, 219.

rondeleti, 200, 246.

typhie, 200, 201, 246.

Siredon, 108,

INDEX.

Sitagra

luteola, 477.

Sitta

cesia, 472.

Smaris, 183.

maurii, 203, 207, 208,

217, 219, 247.

wulgaris, 203, 207, 208,

216, 217, 219, 247.

Smilodon

californicus, 181.

Solaster

endeca, 45.

Somateria

mollissima, 514.

Sorex. 167.

aquaticus, 143.

araneus. 148, 164.

bedfordie, 164,

165.

cristatus, 143,

cylindricauda, 164.

minutus, 164.

quadraticauda, 167.

wardi, 165.

Soriculus

caudatus,

166.

macrurus, 165, 166.

sacratus, 165.

Spathopterus

alexandre, 2.

Spatula

clypeata, 514.

Speotyto

cunicularia, 493.

Spermestes

cucullata, 475.

Tringilloides, 470.

nana, 457.

Spermophila

alboqularis, 478.

cerulescens, 478.

collaria, 478.

eulert, 478.

gutturalis, 478.

hypoleuca, 478.

lineata, 478.

lineola, 478.

nigro-aurantia, 478.

plumbea, 478.

torgueola, 478.

Spermophilus

citillus, 446.

grammurus, 446.

mexicanus, 446.

mongoticus, 446.

tridecimlineatus, 446.

Spermospiza

guttata, 477.

hematina, 477.

165,

Sphagolobus

atratus, 488.

Spheniscus

demersus, 532.

humboldi, 532.

magellanicus, 532.

Sphenocercus

sphenurus, 516.

Sphingurus, 449.

insidiosus, 450.

mexicanus, 450.

prehensilis, 450.

spinosus, 450.

villosus, 450.

Spilogale

putorius, 137, 181.

Spilornis

bacha, 503.

cheela, 503.

spiloyaster, 503.

Spiroptera

uganda, 504, 555.

Spizaétus

bellicosus,

502.

calivatus, 503.

ceylonensts, d03.

coronalus, 2, 503.

. nipalensis, 503.

orientalis, 503.

ornatus, 503.

tyrannus, AUS.

Spizella

socialis, 480.

Squatarola

helvetica, 529.

Starneenas

cyanocephala, 518.

Steatornis

caripensis, 489.

Stenoponia, gen.

391.

celestis, 891, 392.

tripectinata, 391, 392,

393.

Stephanophorus

leucocephalus, 474.

Stercorarius ;

antarcticus, 530.

buffoni, 530.

crepidatus, 5380.

pomatorhinus, 530.

Sterna

cantiaca, 530.

fluviatilis, 78, 530.

hirundo, 78.

minuta, 580.

Strepera

anaphonensis, 488.

Juliginosa, 483.

graculina, 458.

68, 69,

nov.,

INDEX. xvii

Strepsiceros, 349 oles

bustoni, 348, asians Te fp ene

Ens Gee en ay ats D, ve striata, 474.

imberbis, 456 : Sy a 1, a fa et} | Tanagrella

Kudu, 456. y 356. HALES os 353, | — eyanomelena, 474.

Strepsilas Sey eer ee aoe. Ree velia, 474.

interpres, 529 y 195 ic E ee 194, | Tantalus, 322.

Stringops 3 | 203° Bae oe ane loculator, 509.

ataenaiies (oe) 85 Beate ie | Tanyen ae

499. A Se Me Meet Spc

Bh sofiee | foie 193, 218, 219, | megulorhynchus, 498,

flammea, 71, 72, | Synoecns BE ee

492. | australis, 521 | aa nee

perlata, 71, 72. Skea | vmericanus, 453.

punctatissima, 72. aluco. 71. 492 tdicus, 493.

Strongylus an D Neo | _ terrestris, 155.

equinus, 553. GMT Oe ass | Tatu, 124, 141, 149.

Struthidea - i a seme ae Tatusia, Wi 141.

cinerea, 473. a are hybrida, 462.

Struthio, 51, 59 ps a oe kappleri, 462.

aa , ; | senensis, 49”. peba, 462

camelus, 585. | eae es Ge pee

7 } lq 2 7 Se erdianus gu »

gee a nanes 55, | Syrrha ptes Taurus De ce

Sere calis | paradoxus, 519. mevicanus, 154.

defilippi, 481. | ee oe

OF, U7aCU 0.

afin $81, | Tnhyhayin 8,82, nui

contra. 481. aoe Ges, 81, 532. barhara, 138.

Sturnus - | ee 514 Taxidea

menzbieri, 481. | eine ee eal americana, 441.

vulgaris, 48). | ee ti ran Temenuchus

Sula epee ae pagodarum, 481.

Rneermresar. | tae wcus, 479. Testudo

leucogastra, 507. ae 513 elephantina, 2.

piscator, 507. | corn “ide 513. Poleaeces

serrator, 507. Maen) a guacricornis, 455.

Suricata : | eae es ong Tetragnatha

} ar C F “ py =

tetradactyla, 439, | iDance Gt a, 0 3. Serox, 297, 302, 303.

Surni | A — eress@, 300, 3

Way che 493 i leptocephala, 10. | ssn » 300, 301.

a ’ : | Lenioptera ayy Be

Sus cupido, 520.

nengeta, :

PiddaRdnenets AG Te , 484, phen 520.

babyrussa, 140 | peti urogallus, 520.

se Hie 461. ° | Foie 55, 56, 523. Tetraogallus

guineensis, 140. est astute 142 T lamatayensis, 521.

leucomystax, 461 | NEES Genes extor

eee 461, Laat tee ! 43. alector, 477.

orcus, 140 Tanarain ris, 165. Thaumalea

S ra a 5

scr i amherst 7 56

ie ae 461. ; tetradactyla, 133, 463. Ronen Vaal

Sycalis j ; eee ant 446 obscura, 6, ie 8.

arvensis, 479. jaiena idee ae Oi Ss; 5b,

flaveola, 60, 479. | re Tee AC) OE:

PEG, ai ae 149, 446. Thaumasia, 813.

Sylvia oe Sas 169, 384 Thelotornis

o ° j we » 10d, . uP 7

atricapilla, 471. | Tanagra - kirtlandia, 3.

cinerea, 471. 1h ae gras Theristicus

curruca, 471, ae 4 : caudatus, 510.

sylvicola, 471 ee eee ee melanups, 510.

: ; palmarum, 474. Theropithecus

trochilus, 471. sayuca, 474, | gelada, 432

xlviil

Thrasaétus

harpyta, 002.

Thrasops

flavigularis, 181.

Thylacinus

cynocephalus, 100, 466.

Tigriosoma

brasiliense, 508.

leucolophum, 508.

Tinamus

solitarius, 533.

Tinea, 221.

Tinnunculus

dlaudarius, 68, 504.

cenchris, 504.

cenchroides, 504.

dominicensis, 504.

gracilis, \O4.

moluccensis, SOA.

rupicolus, 5O4.

sparvertus, S04.

vespertinus, SOL.

Toccus

erythrorhynchus, 488

melanoleucus, 488.

Tolypeutes

quadricinetus, 141.

tricinctus, 141.

Totanus :

calidris, 529.

Fuscus, 529.

Trachypbarynx,

nov., D2.

nigeri@, 052, 554.

Tragelapbus

angasi, 348.

gen.

buxtoni, 348, 349, 350,

gratus, 456.

scriptus, 2, 456.

sylvaticus, 456.

Tragulus

javanicus, 3, 460.

meminva, 460.

stanleyanus, 4€0.

Tribonyx

mortiert, 920.

ventralis, 3, 526.

Trichechus

manatus, 131.

Trichoglossus

cyanogrammus, 494,

jorstent, 494.

hematodes, 494.

guitchelli, 494.

nove-hollandie, 494.

ornatus, 494.

rubritorques, 4, 494.

Trichosurus

canines, 464.

JSuliginosus, 464.

INDEX.

Trichosurus

orientalis, 464.

vulpecula, 464.

Trigla, 183, 203, 219.

corax, 202.

gurnardus, 203,

210, 217, 219.

hirundo, 202, 216, 247.

Tringa

alpina, 529.

canutus, 529.

Tringoides

hypoleucus, 529.

Triodontophorus, 551.

Trogon, 60.

Trypanosoma

lewisi, 9.

Tupaia

peguand, 444.

tana, 444.

Turacus

buffoni, 66, 491.

corythaix, 491.

livingstoni, 491.

macrorhynchus,

491.

persus, 491.

schalow?, 491.

Turdus

albiventris, 470.

Turnagra ‘

crassirostris, 472.

Turnix, 90, 93.

dussumiert, O24.

lepurana, 524.

sykest, 524.

taigoor, O24.

tankt, 524.

varia, 66, 67, S24.

Turtur

aldabranus, 517.

bitorquatus, 517.

capicola, 517.

cardis, 470.

chinensis, 517.

communis, O17.

falklandicus, 470.

flavipes, 470.

iliacus, 470.

leucomelas, 470.

merula, 470.

migratorius, 60, 470.

musicus, 470.

mustelinus, 470.

erientalis, 517.

pecilopterus, 470.

pilaris, 470.

risorius, 517, 519.

rufiventris, 470.

semitorquatus, 517.

senegalensis, 517.

207,

66,

Turtur

suratensis, 517.

tigrinus, S17.

torquatus, 470.

tristis, 470.

vinaceus, DIT.

wiscivorus, 470.

Tym panistria

bicolor, 518.

Tyrannus

melancholicus, 484.

pipiri, 484.

Uhodon, 297.

hawkesi, 299, 803, 604.

(Zora) frenatum, 306.

Umbrina, 215.

Upupa

epops, 61, 489.

Uria

grylle, 552.

Urobrachya

albonotata, 476.

axillaris, £76.

hocagtt, 476.

Urocissa

flavirostris, 483.

magnirostris, 483.

occipitalis, 483.

sinensis, +83.

Uropsilus, 159.

soricipes, 163.

Ursus, 273.

americanus, 442.

arctos, 139, 442.

— heringianus, 3.

horribilis, 442.

isabellinus, 442.

japonicus, 442.

lotor, 140.

luscus, 139.

malayanus, 442.

muriti ms, +42,

meles, 140.

ornatus, 442.

piscator, 442.

syrtacus, 442.

tibetanus, 442.

Urubitinga

anthracina, 502.

meridionalis, 502.

zonura, 502.

Vampyrus

specorum, 130,

Vanellus

cayennensis, 529.

cristatus, O29.

Vanellus

vulgaris, 529,

Vermipsylla, 365,

Vespertilio

auritus, 131.

discolor, 130, 1381.

leporinus, 131.

moupinensis, 162.

muricola, 162.

murinus, 130, 131.

natterert, 445.

noctula, 445.

perspicillatus, 130.

spasma, 131.

spectrum, 130.

vampyrus, 130.

Vidua

albonotata, 476.

ardens, 476.

paradisea, 476.

principalis, 476.

Vinago

calva, 516.

crassirostris, 316.

delalandii, 516.

Vipera

russelli, 4.

Viverra

civetta, 137, 488.

genetta, 137.

ichneumon, 136.

memphitis, 137.

putorius, 137.

INDEX.

Viverra

tangalunga, 438.

ztbetha, 1387, 438.

Viverricula

malaccensis, 438. °

schlegeli, 438.

Vulpes

sp., 168.

alba, 135.

alopex, 134.

argentatus, 181.

cerulescens, 135.

famelicus, 1.

jilchneri, 168.

hooly, 168.

Vultur

auricularis, 504.

calvus, 504.

monachus, 504.

occipitalis, 504.

Xanthocephalus

icterocephalus, 481.

Xanthomyza

phrygia, 474.

Xanthosomus

flavus, 481.

Xenophrys

monticola, 393, 397,

400, 402.

Xenopsylla

cheopis, 5.

Proc. Zoot, Soc. —1911, No. XX XIX

xlix

Xenorhynchus

australis, 509.

senegalensis, 509.

Xenurus

tricinctus, 462.

unicinctus, 462.

Xerus

erythropus, 446.

getulus, 446,

setosus, 446,

Zapus, 167.

Zenaida

amabilis, 517.

auriculata, 517.

aurita, 517.

Zenaidura

carolinensis, 517.

Zeus, 183, 210, 211, 214,

219

Saber, 209, 216, 248.

Zonotrichia

albicollis, 480.

pileata, 480.

Zorilla

erythrea, |.

Zosterops

dorsalis, 473.

lateralis, 473.

palpebrosus, 473.

simplex, 473.

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EXHIBITIONS AND NOTICES. 5

age

Tt Sucrerary. Report on the Additions to the Society's Menagerie during the months of

June, July, August, September, and October, 1910 ...... 2... cece cece reese ce eees il

Mr. RB. I. Pocock, F.LS., F.Z.8. xhibition of a living specimen of the Black Rat (Mus

ratiws) from Sark............ bia lech sled seinen (Nebel a teiete AU CEN Eon cored OC Leaowodds 4

Mr. R. I. Pocock, F.L.S., F.Z.8. Exhibition of a female hybrid between a male Black

Lemur (Lemur ee) and a female of the Red-fronted variety of the Fulvous Lemur

(Lemar frlos rufifrons) «..s2.cerceesseees AG ciaeuren as. oa dbeC ace in uaceelaCotar everett 5

The Hon. N. Cuartes Rornscurp, M.A., F.Z.8. Exhibition of some Fleas interesting in

connection with the spreading of plague by these insects ......0+...2.. seer es ae yt

Mr. J. Luwis Bonnorz, M.A., F.L.S., F.Z.8. Exhibition of, and remarks upon, a young Cairo

Spiny Mouse (Acomys cahir ns TOMAS epee Uaioe ovatate lar eltanscn te aces See AO Mie sia c 5

Mr. J. Lewis Bonnors, M.A., F.L.S., F.Z.S. Exhibition of, and remarks upon, a pair of

hybrids between the ratible Hitch (Fringilla montifringilla) Zine the Chaffineh (Frin-

GUM COHICUS) totam sn «sis 'aleelnle\e to PAN ae tS te SA ah Nn pn ee dane STN Na Pee ON

Mrs. R. Hate Tuomas, F.Z.8S. Exhibition of, and remarks upon, a series of skins llus-

trating an experiment in Pheasant-breeding ......--....-- EMER BN DOG bc uc 6

Dr. W. Nicott and Prof. E. A. Mincaty, M.A., V.P.Z.S. Exhibition of, and remarks upon,

two species of Cysticercoids from the Rat-flea (Ceratophyllus fasciatus). (Text-figs.1&2.) 9

Dr. Wrextam Nicouu. Demonstration of his method for the collection of Trematodes .... 45

Dr. R. ©, Lereer, F.Z.8. Exhibition of the Nematode infection known as Onchocerciasis.. 40

“

Dr. J. F. Gunmitz, M.A., D.Sc. Notice of a paper on “ The Development of Solaster endeca

Homes ai Actes aatte ta ave) Rais SS ave Ba ete orcoiete(tss 6 alee shelter eneale hypo nl can eUameinvs te ete Sis sins 45

Mr. D. Sern-Smirn, F.Z.8. Exhibition of, and remarks upon, living examples of the

Australian Budgerigar or Undulated Grass-Parrakeet (Melopsittacus undulatus),

showing three colour-phases ......++..-0+.+4+ RR Siena AO, ese 2 oe cha etbies uated

The Srcrerary. Report on the Additions .to the Suciety’s Menagerie during the month

Of Novem berlOl ayer aersiees cee aie teneyeretotade tae iayailalapuiecapensnaie 's0s here foreyetal azo Net oa 100

Dr. H. Hawmonp Siti, M.R.OS., F.Z.S8. Exhibition of, and remarks upon, a specimen

of the Red Grouse, Lagopus scoticus, which displayed a curious variety of the ordinary

PUM are eal arvana aya areata ome ee TRG, CA A CaN er ete Beata crtrim nate od sis se OO)

Contents continued on page 8 of Wrapper.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

EXHIBITIONS AND NOTICES.

November 15, 1910.

Dr. S. F. Harmer, M.A., F.R.S., Vice-President,

in the Chair.

THE SecRETARY read the following report on the additions made

to the Society’s Menagerie during the months of June, July,

August, September, and October, 1910 :—

JUNE.

The registered additions to the Society’s Menagerie during

the month of June were 405 in number. Of these 254 were

acquired by presentation, 39 by purchase, 18 were received on

deposit, 12 in exchange, and 82 were born in the Gardens.

The total number of departures during the month, by death

and removals, was 142.

Amongst the additions special attention may be called to the

following :—

2 Addra Gazelles (Gazella ruficollis) 3 2 , new to the Collection,

1 Mountain Oribi (Owrebia montana), 1 Sabre-horned Oryx (Oryx

algazel) Q, 1 Addax (Addax nasomaculatus), 1 Soudan Fennec

(Vulpes famelicus), 2 Erythrean Zorillas (Zorilla erythrea),

1 Dongolan Genet (Genetta dongolana), and 2 Banded Mongooses

(Crossarchus fasciatus), from the Soudan, presented by Gilbert

Blaine, Esq., F.Z.S., on June 4th.

Proc. Zoou, Soc.—1911, No. I. 1

THE SECRETARY ON ADDITIONS TO THE MENAGERIE,

1 Addra Gazelle (Gazella rujficollis) 3, new to the Collection,

from the Soudan, presented by G. Guy Chetwynd, Esq., F.Z.S., on

June 4th.

1 White-spotted Squirrel (Fwnisciwrus leucosiagma) and

1 Speckled Squirrel (Scturus punctatus), both new to the Collec-

tion, 1 Gaboon Viper. (Bitis gabonica), and 1 Nose-horned Viper

(Bitis nasicornis), from Dunkwa, Gold Coast, presented by Dr. H.

G. F. Spurrell on June 18th.

6 Cocks-of-the-Rock (Rupicola:crocea), from British Guiana,

deposited on June 14th.

1 Red Bird-of-Paradise (Paradisea rubra), presented by

J. M. Doctor, Esq., on June 11th.

1 Hunstein’s Bird-of-Paradise (Diphyllodes hunsteins), received

in exchange on June 23rd.

] Short-tailed Parrot (Pachynus brachyurus), from the Upper

Amazons, presented by H. D. Astley, Esq., F.Z.8., on June 14th.

2 Queen Alexandra’s Parrakeets (Spathopterus alexandre),

from Western Australia, purchased on June llth. .

2 Condors (Sazcor hamphus gryphus), from Chili, pr esented by

the Government of Chili on June 27th.

A collection of Birds, including Tanagers, Sugar-birds,

Tinamous, and Conures, from Brazil and Chili, presented by

Albert Pam, Esq., F.Z.S., on June 18th and 27th.

A collection of Birds from Venezuela, including various

Tanagers, Finches, a Naked-eyed Pigeon (Columba gymno-

phthalma), and an Ori inoco Goose (Chenalopex jubatus), presented

by Hugo Pam, Esq., on June 21st.

5 Elephantine Tortoises (Testudo elephantina), from Aidabra,

presented by Edward Mayer, Esq., on June 15th.

JULY

The registered additions to the nee s Menagerie during the

month of July were 327 in number. Of these 124 were acquired

by presentation, 16 by purchase, 85 were received on deposit,

15 in exchange, and 87 were born in the Gardens.

‘The total number of departures during the month, by death

and removals, was 189.

Amongst the neetblons special attention may be called to the

following :—

1 Bufton’s Kob (ikebas kob), from Sierra Leone, presented by

J. A. Tinling, Esq., on July 8th.

- | Sing-Sing Waterbuck (Kobus unctuosus) and two Telecel

Bushbucks (Zragelaphus scriptus), from Gambia, presented by

Cape: Sir George Denton, K.C.M.G., F.Z.8., on J uly 19th.

- 2 Harnessed Bushbucks (Tragelaphus scriptus), 1 Black-tailed

Or ab) (Ouredia nigricaudata), and 1 Crowned Hawk-EHagle

(Spizaétus coronatus), from French Guiana, presented by

Fenwick Owen, Hsq., on July 19th.

1 Abyssinian Duiker (Cephalophus abyssinicus), from Nigeria,

presented by W. A. Clayton, Esq., on July 20th.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 3

2 Black-backed Geese (Sarcidiornis melanonota), from India,

presented by Lt.-Col. D. C. Phillott, on July 25th.

3 Wattled Peewits (Lobivanellus lobatus), 3 Black-tailed Water-

hens (Zrihonyx ventrahs), and 1 Tufted Umbr re (Scopus umbretta),

hatched and reared in foe Menagerie.

AUGUST.

The registered additions to the Society’s Menagerie during the

month of August were 200 in number. Of these 105 were

acquired by presentation, 15 by purchase, 27 were received on

deposit, 29 in exchange, and 24 were born in the Gardens,

The total number of departures during the month, by death

and removals, was 181.

Amongst the additions special attention may be ealled to the

following :—

3 Grey Crab-eating Dogs (Canis griseus), from the Argentina,

presented by Wilfred Smithers, Esq., on August 22nd.

1 Korin Gazelle (Gazella rufifrons) 2 , from Senegal, presented

by Maxwell Lyte, Esq., on August 2nd,

2 Ross’s Plantain-eaters (Musophaga rossce) and 1 Ardesian

Hawk (Cerchneis ardesiacus), both species new. to the Collection,

from Uganda, presented by L. M. Seth-Smith, Esy., on

August 15th.

3 Pigmy Cormorants (Phalacrocorax pygmeus), from Dobrud-

scha, Roumania, presented by the Hon. N. C. Rothschild, F.Z.8.,

on August 25th.

A collection of Birds, including 2 Rough-billed Pelicans

(Pelecanus trachyrhynchus), 2 Roseate Spoonbills (Ajaja rosea),

and 3 American Darters (Plotus anhinga), received in exchange

from the Zoological Society of Washington on August 23rd.

1 Green Tree-Snake (Dendraspis viridis), 1 Kirtland’s Tree-

Snake (Thelotornis kirtlandiz), and 1 Banded Gecko (Hemidactylus

fasciatus), all new to the Collection, from the Gold Coast,

presented by Dr. H. G. F. Spurrell on August 21st.

SEPTEMBER,

The registered additions to the Society’s Menagerie during the

month of September were 228 in number. Of these 3/7 were

acquired by presentation, 81 by purchase, 47 were received on

deposit, 53 in exchange, and 10 were born in the Gardens.

The total number of departures during the month, by death

and removals, was 210.

Amongst the additions special attention may be called to the

following :—

1 Brown Bear (Ursus arctos beringianus), from Kamschatka,

presented by W. 8. Race, Esq., on Sept. 30th.

4 Javan Mouse-Deer (Tragubus 4 jovanicus), from Java, purchased

"on Sept. 26th.

4 MR. R. I. POCOCK ON THE BLACK RAT.

1 White-naped Roller (Coracias navius), from the Gambia,

new to the Collection, deposited on Sept. 15th.

1 Red-collared Lorikeet (Trichoglossus rubritorques), bred in the

Menagerie.

1 White-winged Wood-Duck (Asarcornis scutulatus), from

Burma, new to the Collection, presented by Major H. H.

Harington, F.Z.S., on Sept. 12th.

A collection of Snakes containing, amongst others, 6 Indian

Cobras (Vaia tripudians), 2 Banded Kraits (Bungarus fasciatus),

and 4 Russell’s Vipers (Vipera russelli), received in exchange on

Sept. 10th.

OCTOBER.

The registered additions to the Society’s Menagerie during the

month of October were 202 in number. Of these 102 were

acquired by presentation, 26 by purchase, 47 were received on

deposit, 21 in exchange, and 6 were born m the Gardens.

The total number of departures during the month, by death

and removals, was 225.

Amongst the additions special attention may be called to the

following :—

1 Siamang Gibbon (Symphalangus syndactylus), from Sumatra,

deposited on Oct. 26th.

1 African Cheetah (Cyncelurus jubatus), from Andamowa,

German Cameroons, presented by Walter Watts, Hsq., on

Oct. 22nd.

2 Indian Buffaloes (Bubalus buffelus), from India, received in

exchange on Oct. 15th.

2 Giant Bustards (Hupodotis kort), from P. W. J. Le Roux, Esq.,

a Black-breasted Harrier-Eagle (Circaétus pectoralis), from

Crossland Robinson, Esq., and a Stanley Crane (dAnthropoides

paradisea), from Dr. Robert Broom, C.M.Z.S., all presented to

the King’s South-African Collection through Dr. L. Péringuey,

F.Z.S., Secretary of the South-African Presentation Committee,

and deposited in the Gardens on Oct. Ist.

A collection of Birds from Venezuela, presented by Hugo

Pam, Esq., on October 11th, containing a Bare-faced Hangnest

(Gymnomystax melanicterus), a Chimachima Milvago (Milvago

chimachima), a Violaceous Night-Heron (Vycticorax violaceus),

3 Naked-eyed Pigeons (Columba gymnophthalma), and others,

5 Jackson’s Whydah-birds (Drepanoplectes jacksoni) and

2 Crimson-ringed Whydah-birds (Pentheiria laticauda), from

East Africa, new to the Collection, presented by Mis. George Style

on Oct. 22nd.

Mr. R. I. Pocock, F.L.S., F.Z.8., exhibited a living specimen

of the Black Rat (Jus rattws) from Sark, which had recently

been presented to the Society’s Menagerie by Mrs. C. Russell.

ON NEWLY-BORN CAIRO SPINY MICE, 0

Mr. R. I. Pocock also exhibited a female hybrid, bred in the

Society’s Gardens, between a male Black Lemur (Lemur macaco)

and a female of the Red-fronted variety of the Fulvous Lemur

(Lemur fulvus rujifrons), and pointed out that the offspring re-

sembled neither of its parents. It had not inherited the facial

fringe of its father nor the white over the eyes and on the fore-

head of its mother, this avea of the head being dark ashy black

with a deeper tinted central line, more approaching the colour

seen in the Black-fronted variety of the Fulvous Lemur (ZL. fulvus

nigrifrons).

The Hon. N. Cuarntes Rotuscuiyp, M.A., F.Z.8., exhibited the

following species of Fleas, which were of interest in connection

with the spreading of plague by these insects :—Pulex irritans,

AXenopsylla cheopis, Ctenocephalus canis, C. felis, Ceratophyllus

Jasciatus, C. londiniensis, Clenopsylla mayardt.

Mr. J. Lewis Bonuors, M.A., F.L.S8., F.Z.8., exhibited in

spirit a young Cairo Spiny Mouse (dAcomys cahirinus) about 12

hours old, and pointed out the advanced state of the young at

birth as compared with the common House-Mouse and other

species of the genus Jus.

In Acomys the young at birth were of a pale slate-colour and

sparsely covered on the upper parts with downy hairs 2 to 3 mm.

in length. Across the lower part of the back the spines could

be distinctly felt.

In spite of this advanced development the period of gestation

was only 11 days at the most. In the case under notice the male

was introduced at 7 p.m. on 19th July and the young were born

between midnight and 9 a.m. on the 3lst July, a maximum

period of 11 days and 14 hours. The period in the House-Mouse

was about 13 days. In Acomys the usual number in a litter was

three ; four were exceptional, but two were not uncommon.

In correlation with their forward development the young were

also very large at birth, as a comparison of measurements of the

individual exhibited and the average measurements of an adult

male show :—

Young

12 hours old. Ad. g.

Head and body .... 51 mm. 101 mm. about 3 full size.

“| Bo aa a aa eit 35 mm. 105 mm. “ai a ek

ind: foot)... 5. 14 mm, 18 mm. ieee ae

late sss ore cies siecicitis ~ ) so enum 17 mm. pel ee ae

When 8 to 10 days old the young were able to take care

of themselves, and on the fourth day after birth their eyes were

open and they occasionally left the nest and ran about the cage.

They bred at a very early age, and by the agency of a young male

born on the 4th June progeny were produced on the 3lst July,

so that it must have bred when only 46 days old.

6 MRS. R. HAIG THOMAS ON

Mr. Bonhote further stated that among Sand-Rats of the genus

Meriones, a considerable number of which he had bred this - year,

the period of gestation was at least 17 days, and the young, which

were born quite naked, looked after themselves in about three

weeks, five being the usual number in a litter.

Mr. J. Lewis Bonnore, M.A., F.L.S., F.Z.S., exhibited a pair of

hybrids between the Bramble-Finch (Fringilla montifringilla) and

the Chaffinch (fringilla celebs). 'This cross had been bred for the

first time in 1907 by a lady in Kent, who had since bred several

every year.

Mr. Allen Silver, a well-known fancier, first called Mr. Bonhote’s

attention to these birds. He wrote:—“The young in nestling

plumage chiefly resemble a nestling Chaflinch, but sy traces of

the white rump.

“Males from the cross in which the Brambling was the cock

are in every case more brilliant and richer in tone, and exhibit a

marked Bramble-Finch form of head, more so than the cross in

which the Chaffinch was the male parent. The heads of the

males in this latter cross (Chaffinch g and Brambling ©) are less

speckled and almost even in tone of colour, showing few signs of

the dark bases to the feathers on the head and having more of a

Chaffinch shape.”

The cock exhibited resembled a Bramble-Finch on the head,

rump, tail, and vent, but on the mantle and breast the colour

showed a mixture of the two species ; the tail also showed more

white than was usually found in a pure-bred Bramble-Finch.

A peculiar feature of this bird was a slight “ peak ” crest, which

had not appeared in any of the other specimens. It seemed to be

caused by the long feathers of the nape (a Bramble-Finch feature)

meeting the long feathers of the crown (a Chaftinch feature).

One other male of this cross that Mr. Bonhote had seen closely

resembled a hen Bramble-Finch.

The hen bird exhibited showed the Chaftinch blood more

clearly, her only Bramble-Finch features being the two parallel

stripes on the nape, the partially white rump, traces of yellow on

the wing-bars, and a more yellowish tint on the breast. The

fertility of the hybrids had not yet been tested.

Mrs. R. Hate Tuomas, F.Z.S., exhibited a series of skins illus-

trating an experiment in Pheasant - breeding, and gave the

following account of the production of Thawmalea obscura in the

F, generation from a cross between Thawmalea amhersti 2 and

T. picta 3 —

“The skins of the birds used in this experiment are exhibited

together with their pedigree (see page 7). An Amherst cock is

placed with these skins to show the plumage, but the bird took

no part in the experiment. ;

ASANT-BREEDING,

NIAL PHI

EXPERIMI

Purchased from Jamrach, 1905, one Thaumalea amhersti 9. Purchased from Col. Middleton, The Chase, Herefordshire,

Purchased from Mrs. Adair Campbell, Ardleith, Cardross, 1906, one Thaumalea picta g.

1906, two Thaumalea amhersti 2.

Mating 1907.

3 Thaumalea amhersti 9 X Thaumalea picta 6.

Dygees tis | | | | | ee | |

9 g 9 2 3 3 3 3 é é

Matings 1909 in tivo pens.

Cream | Russet | dark Brown (obscura)

Thaumalea obscura Fy dead chicks 1I—~> 4 5 2(9)

produced in I’, F;2 XF,¢ | 49 eggs, 29 infertile, 20 hatched | F) 9 XFid

from both pens. ; : : . :

9 of these Fy birds reared. 17 of these Fy birds alive 1910.

F; | | | | | | | |

? 4 ? z dg $ g (obscura)

Matings 1910 in two pens.

Fo? XFod (semi-obsewra). F.2 X Fog (obscura).

Ee eee Voy tee oe | | | | Soa ae ae

Se g ) 9 9 9 é 3 3

(cream) (cream) (obscura) (obscura) (obscura) (obscura) (cream) (cream)

All 8 chicks lived only a few days.

dark Brown (obscura)

Summary of F3 Colours { Cream

1910 4

8 ON EXPERIMENTAL PHEASANT-BREEDING.

“In Elliot’s ‘ Phasianide’ is a coloured plate of a pair of

Thaumalea obscura with their young, which were considered by

him and some other ornithologists as a variety of picta; according

to this plate the birds apparently bred true.

“In my pheasantry in 1907, three Amherst hens were mated

with a Golden cock and produced a number of young called F, in

the pedigree. In 1909 two pairs of these F, birds were mated,

and from these two pens 20 F, chicks were hatched. Amongst

them were three chicks of a deep chocolate-brown, in startling

contrast to the others, which were cream-colour with a bronzing of

russet on the back and throat. The down of these obscura chicks

was of a uniform dark brown to the skin, and each eye was

rimmed with a finely pencilled cream line, which was connected

by another cream line across the top of the bill, giving the quaint

appearance of a pair of spectacles; there was also a cream patch

on the throat, varying in size in each individual, sometimes being

a mere spot. The legs were a dull olive-brown. These brown

chicks were produced from both pens, and were of both sexes,

but unfortunately only one of the three (a cock) was reared.

Thaumalea birds attain adult plumage only in the second year, so

it was not till July this year that my surmise that obscura

had been produced was confirmed. The cock has the same barred

tail (pattern Amherst, coloration Golden), and the same dark

brown on head and throat and neck, as the bird depicted on Elliot’s

plate, but his breast is a duskier red and the crest and mantle

paler. When a chick, his coloration was similar to the young

shown on Elliot’s plate, only the brown was a deeper, richer shade.

This F, cock was mated in the spring of 1910 with two F, hens,

his sisters. The darkest hens were picked out, with Golden eye-

skin (yellow with red round the rim); Amher st hens are lighter

and greyer in plumage than the Golden, and the eye-skin is a

greenish blue. From this mating eight birds were hatched, four

cream-coloured and four dark brown chicks; these last had pre-

cisely the same coloration as that of the father. The numbers

tally with those to be expected according to Mendel’s law. I

hope to obtain more evidence next season from the same birds,

and to have better luck, for all these eight chicks lived only a few

days. As I intend to breed again from him, I cannot show you

the skin of the adult obscaa cock, but only some feathers plucked

from his breast and back, and the skins of his two brothers and

of two of his sisters. An examination of these breast-feathers

plucked from obscura and of some of the breast-feathers of the

cock marked ‘ semi-obscwra’ will show the reason of the so-called

‘duskiness’ of hue; each feather has a bar of metallic green on

the inner half, while the outer end of it is red. You will observe

in the Golden cock that the breast-feather is red throughout.

“ Apparently, then, Zhawmalea obscura is a hybrid recessive

mutation, breeding true, as shewn in Hiliot’s plate and also in my

experiment this year, according to Mendel’s law. I have used

the word ‘mutation,’ but these birds may possibly be a reversion

to the ancestral parent form of both “ species” of Zawmalea.

ON TWO SPECIES OF CYSTICERCOIDS, 9

“By whatever name it is called, this hybrid is undoubtedly a

homozygote for pattern and colour, pure and permanent, trans-

mitting these characters to its descendants. Cross-breeding

between these two varieties of Thawmalea produces a new form,

owing possibly to the meeting of characters never previously com-

bined, and as they are constant it is evident these have an affinity

and have become inseparable.

“On such lines Evolution might be conceived as having pro-

ceeded fairly rapidly towards the separation of species.

“Some while ago 1 showed a hybrid hen whose pure Swinhoe

plumage had been transmitted through the cock. To-day the

skin of her son, a pure Swinhoe cock, is exhibited for your

inspection.

** An account of my method and management of these Pheasant-

breeding experiments has been already published in the ‘ Pro-

ceedings ’ (1909, p. 885), showing the care and precautions used.”

Iwo Species of Cysticercoids from the Rat-flea (Cevatophyllus

fasciatus).

(Text-figures 1 & 2.)

Dr. W. Nicoxt and Professor EK. A. Mincuin, M.A., V.P.Z.S.,

exhibited examples of two species of Cysticercoids which had

been found in the body-cavity of the rat-flea (Ceratophyllus

Jusciatus) dissected by Professor Minchin in the course of in-

vestigations upon the development of Zrypanosoma lewisi in the

flea.

The first specimen shown (text-fig. 1) was one previously ex-

hibited by Prof. Minchin (P. Z. 8. 1909, p. 741). Since then

experiments had been carried on by Dr. Nicoll at the Lister

Institute, which proved that it was the larval form of Hymenolepis

diminuta, and threw light upon the mode of infection. This

Cysticercoid had occurred in about four per cent. (8 in 207) of all

the fleas examined during a period of thirteen months. It was

found in the body-cavity and usually only one specimen at a

time. As many as three, however, had been found in one flea.

In the retracted state the Cysticercoid consisted of a flattened

oval body and a more or less elongated tail. The dimensions of the

body were 31 xX ‘25 mm., and the tail might be as much as ‘8 mm.

long. The wall of the body consisted of a number of well-

differentiated layers. The outermost layer was non-cellular and

bore a certain resemblance to cuticle. It was marked by fine

radial striations. |The second layer consisted of columnar cells,

each with a large and distinct nucleus. Neither of these layers

occurred in the tail. The third layer was parenchymatous,

consisting of a large number of irregularly disposed cells. This

was continuous with the substance of the tail. The next layer

was apparently of a fibrous nature, consisting of a few cells set

in a mass of circularly arranged fibres. This layer was separated

from the next by an intervening space. This was seen usually

10 DR. W NICOLL AND PROF, E. A. MINCHIN ON

only as a narrow fissure. It represented the very much reduced

vesicle, and separated the main part of the body from the part

which was eventually evaginated to form the head of the tape-

worm. Internal to this was another layer of loose parenchyma,

which at the posterior pole merged into the tissue of the scolex.

The latter was extremely small (075 x-09 mm.). Onit there were

four small circular suckers (diam. ‘055 mm.) and a small unarmed

rostellum. .

Cysticercoid of Hymenolepis diminuta. X 125.

This description tallied almost exactly with that of Grassi and

Rovelli (Atti Acc. Gioenia Sc. Nat. Catania, iv. 1892, pp. 31-33,

pl. iv. fig. 3). They found a corresponding Cysticercoid in the

beetles Akis spinosa and Scaurus striatus, in Anisolabis annulipes

and in the larva of Asopia farinalis. By feeding experiments

with Cysticercoids derived from the first of these, they were able

to produce infection in a man, and the tapeworm which was

recovered was Hymenolepis diminuta Rud. (= Tenia leptocephala

Creplin). *

It was recognized that the Cysticercoid from the rat-flea was

very probably the larva of Hymenolepis diminuta, but that could

not be definitely stated without further information, for two

other rat-tapeworms (H. relicta Zschokke and H. horrida

v. Linstow) are known, closely resembling H. diminuta and

probably possessing larvee almost indistinguishable from that of

H. diminuta. On that account, in order to establish the identity

of the Cysticercoid, a series of feeding experiments was undertaken.

TWO SPECIES OF CYSTICERCOIDS. 11

For this purpose a litter of young rats was separated from their

mother as early as possible and isolated. They were fed entirely

on boiled bread and milk. They were kept under observation

for three months, during which time frequent examination of

their feeces showed that they were free from tapeworms. Two of

the rats were then fed daily with the fleas, which were mashed up

in their food. This was continued for 19 days, feeding being

omitted on three of these, when one of the rats died. During

this period the feces were examined regularly, but no ova were

found. On examining the dead rat, five specimens of Hymenolepis

diminuta were found in the intestine. They were of various

sizes, the largest being over 40 cm. long and the smallest only

lem. Next day numerous ova were found in the feeces of the

other rat. From this it was evident that the complete develop-

ment of Hymenolepis diminuta took place in less than three

weeks. Grassi and Rovelli found ova after 15 days, which was

probably about the actual length of time required. The remaining

rats of the litter were kept under observation for a further period

of two months, but during that time no ova were found in the

feeces of any of them. The same experiment was repeated later

with a similar result, ova appearing in the feeces on the 19th day.

The rat which survived the first experiment was eventually

killed and nine tapeworms were removed from its intestine. <A

total of 14 tapeworms was therefore produced from the 340 fleas

with which the rats had been fed. This indicates that about

four per cent. of the fleas contained Cysticercoids, and the result

agrees with the number of Cysticercoids found in fleas which were

actually dissected.

The supposition, put forward in the previous communication,

that the flea becomes infected in its larval state, had to some

extent been confirmed. The intact ova of Hymenolepis diminuta

were found in the intestines of 2 larve out of 145 examined. No

stages in the development of the Cysticercoid, however, had been

yet met with in the larval flea. On the other hand, a very young

Cysticercoid had been found in the body-cavity of one out of a

dozen pupz examined. The presumption was that it was only

the oldest and largest flea-larve that were capable of ingesting

the tapeworm eggs and that no development took place until the

beginning of the pupal stage.

The second Cysticercoid (text-fig. 2, p. 12) was of much greater

interest than the first. Hitherto only a single specimen had

been met with, and it occurred in the body-cavity of a male

Ceratophyllus fasciatus. It was considerably smaller than the

first Cysticercoid, and like it consisted of an oval body and a tail.

The former was °16 mm. long and the latter 19mm. The two

outermost layers of the body-wall were the same as_ before,

but the parenchymatous and fibrous layers were not so sharply

differentiated from each other. The scolex, again, was very much

larger (diam. ‘096 mm.) and it had a rostellum armed with a

single row of spines,-23:in number. ‘The length of each spine was

12 DR. W. NICOLL AND PROF, E. A. MINCHIN ON

‘017 mm., the handle being ‘01 mm. and the guard :007 mm.

long. The prong was of the same length as the guard. There

were four well-formed suckers, which had been pressed into a

somewhat crescentic shape (greatest diameter -042 mm.).

Text-fig. 2.

Cysticercoid of Hymenolepis murina. X 250.

a. Rostellar spine in full lateral view. XX 600.

b. The same, somewhat tilted.

Besides the two Cysticercoids described here and the well-known

Cysticercoid of Dipylidium caninum from the dog-flea and human

flea, there was only one other instance on record of a Cysticercoid

being found ina flea. This observation was made a few months

ago by Dampf (Centralbl. f. Bakt. ete. lte Abth. Orig. liv.

pp. 452-4), who found in a new species of flea Wesopsylla eucta from

the jerboa (Alactaya jaculus) a Cysticercoid bearing a close resem-

blance to the second of those described here. Dampf was obliged

to make his examination from a mounted (type) specimen of the

flea, and on that account could not be certain of some of the

details, as, for instance, the exact number of the spines and the

presence of a tail. So far as he could make out, there were from

bel

TWO SPECIES OF CYSTICERCOIDS,

20 to 24 spines. Lihe, to whom the specimen was referred,

mentioned the supposition that the Cysticercoid might be that of

Hymenolepis nana (v. Sieb.),

From the description given of the second Cysticercoid it was

apparent that it was very like the scolex of Hymenolepis murina

Duj., a frequent parasite of various species of rats. The size of

the scolex of the Cysticercoid, the arrangement, number, size, and

shape of the spines all agreed with those of Hf ymenolepis murind,

and it was certainly the only rat-tapeworm to which the Cysti-

cercoid could be referred. The closely allied Hymenolepis micro-

stoma Duj. had 30 spines of much smaller size. HH. contracta

Janicki and H. muris variegati Janicki, neither of which corre-

sponded to the Cysticercoid, were the only other armed Hymeno-

lepids found in rats, and the authors were forced to the conclusion

that this Cysticercoid must either be referred to H. murina or

to some other as yet undescribed form, the scolex of which was

indistinguishable from that of Z. murina.

Hitherto the lite-history of Z. murina had been believed to be

of a unique type. According to the researches of Grassi and

Rovelli (op. ev. pp. 75-83) this tapeworm dispensed altogether

with an intermediate host, and they had demonstrated that it

passed its encysted stage in the walls of the intestine of the rat

itself. This was supported on general grounds by the fact that

this particular tapeworm usually occurred in enormous numbers,

which would involve the ingestion of a correspondingly large

number of intermediate hosts. The matter gained interest from

the fact that H. murina was believed by some to be identical with

H. nana, a dangerous tapeworm of man, although the identity was

still a matter of dispute. In size and structure the two species

were indistinguishable, and H. nana like H. murina occurred in

enormous numbers. Further, no intermediate host had yet been

discovered for H. nana. Now the discovery in the rat-flea of this

Cysticercoid bearing such a close resemblance to the scolex of

H. murina \ed to one of two conclusions: either that, as already

mentioned, there might be some undescribed tapeworm of which

the Cysticercoid was the larval stage, or that the rat-flea might

function as an intermediate host of H. murina. The latter

supposition, even if correct, did not necessarily disprove Grassi and

Rovelli’s results. Rats certainly ingested a large number of fleas

from time to time, but it was difficult to imagine that they would

acquire a very large infection with H. murina by this means;

and if H. nana be identical with H. murina it was scarcely

conceivable that a human being could accidentally swallow a

sufficient number of fleas to give rise to an infection of over a

thousand tapeworms, as was frequently found to be the case. It

would still therefore be necessary to admit that the development

of H. murinaw usually took place without an intermediate host,

but that occasionally the rat-flea might function as such. With

such a conclusion the life-history of H. murina became even

more remarkable than before.

14 MR. J. LEWIS BONHOTE ON THE

PAPERS.

1. On the Inheritance of the Webfoot Character in Pigeons.

By J. Lewis Bonyorn, M.A., F.L.S., F.Z.8.

[Received August 29, 1910: Read November 15, 1910.]

(Text-figures 3 & 4.)

In the P. Z.8. for 1905, p. 550, Mr. Staples Browne published

a paper on the webbed-foot character in Pigeons, and from a

series of careful experiments conducted by him the results seemed

to show fairly conclusively that the webbed foot was a Mendelian

character and was recessive to the normal or non-webbed foot.

In this paper the author drew an arbitrary line and counted as

webbed all birds which showed on one foot a web at least to the

first interphalangeal joint of the 2nd and 38rd digits and to the

second interphalangeal joint of the 4th digit, any bird with

less webbing being considered as normal. In my experiments

T have followed his distinction for the sake of uniformity. All

his results, with one exception, go to show that the webbed foot

behaves as a pure Mendelian recessive character.

In the aberrant case two F, birds were mated together for two

seasons, producing altogether 23 birds, in none of which the web

appeared. Both these F, birds were subsequently tested and proved

to be carrying the webbed character.

In the spring of 1905, Mr. Staples Browne kindly gave me a

pair of his webbed birds [Exp. 14, Joc. cit.], and during that year

and the two subsequent years this pair produced nothing but

webbed birds.

- In 1907 Mr. F. W. Smalley, a well-known and successful breeder

of fancy Pigeons, wrote to me in regard to some webbed birds

which tended to make their appearance from time to time in his

Taste I, r

Exp. 9 Origin| Also Origin} Also | Nature of Feet.

WeNG(G) Eee. e N (126)| 6 ae DRXDR 6 2

4) IST (BD) | coe “a N (426)| a at D XDR 6 0

| N (61) ats a N (104) oes ae DRXDR 6 2

6 i

| N10) | 2 u ie (01)| .. | DRXDR

stud. At my request he kindly sent me their pedigrees, from

which it will be seen that the inheritance was strictly Mendelian

and entirely bore out Mr. Staples Browne's results. .

WEBFOOT CHARACTER IN PIGEONS. 15

Text-fig. 3.

No x No NéxNo

EXPAN C. Exp} Oo.

SoTHERSN 2W No 6N 2w N 35 OTHERSN

W69 .2? 4N

Pedigree of Mr. Smalley’s bird (W 69).

(Birds underlined are supposed heterozygotes.)

A further and striking confirmation of this came to light this

year (1910), when a web-footed bird suddenly appeared in whose

direct ancestry on either side no webbed birds had been known

for at least six generations; an investigation of the pedigree,

however, showed one of the birds to have been a possible hetero-

zygote, and on this assumption, and presuming all the subsequent

matings to have been DR x D, the case is quitesimple. To some

Tasxe IT.

Exp. |, 9 Origin) | Also 3 Origin) Also | Nature of eet

e N (5) a fo ING(60)) |e: ee DRXD 3 0

if N (52) € $8 ING (80) eee ia DRXD 7 0

g INK(72)8 | Perey NCL) sees i DRxXD 4 0

he le Ni(46) | 8 a TN OD gf S69 DeXDR | 15 0

ee NT (4876)) oe (a INR SBAN De x<DRi) 13 0

k ...N (5486)} i Ea ING@eERIN Ae | DRDRi! 9 1

it- may appear strange that a DR bird should have been chosen

in so. many successive years, when the chances were even on the

bird being.a DB..or pure Normal.. It must, however, be re-

membered that since young were.reared from a large proportion

16 MR. J. LEWIS BONIIOTE ON THE

of the birds noted in Table II., one would, in theory, have

expected the web to have made its appearance in more than one

instance, as the number of DR birds in Mr. Smalley’s lofts must

be considerable.

The fact remains that a webbed bird has been bred, and if its

pedigree were not well-known one would have had to regard it

as a “sport,” without being able to account for its presence in

any way.

So far, therefore, from well-attested evidence of independent

breeders, the webbed character seems to be established beyond

doubt as a Mendelian recessive.

In the summer of 1907, however, Mr. Smalley sent me a male

web-footed bird (W 69), which I paired at once with one of the

progeny (also webbed) of the pair of birds originally given me by

Mr. Staples Browne. My object was merely to get a change

of blood and to keep up a small stock of webbed birds.

Both the parents were webbed and, as we have seen, their

ancestry showed that in both families the web had acted as a

Mendelian recessive, so no doubt was felt but that the whole of

the progeny would also be webbed. The result of this mating

was five birds, of which four were normal.

For the past three years I have carefully mated these five

birds and their descendants, with the results as shown in the

following table :—

Taare if.

-. | Also . - | Also | Nature Feet.

Exp. Origin Origin :

2. from | Used 3+ | from mane M of | Normal.| Webbed

in In Mating.

ee ee ee |— — an

lw (7)| 14S8B Wi(69))| td | eR IR ee 1 ; Mixed

|W (a) ge 9) adee| Noa) ar alae ante 1 6: iG) sas,

|N (b) 1 4 || N a) Ty ete 1 1

N (8) 1 Sripseiayih the le we 5 3 ;

Ww (24)| 2 | 12 || W (9)|148B ig 1 0 ; oe

N (29) 4 W (35) 2 In 5 1 aT ' ;

N (22) 4 | N (19) 4 ; theory Gaye] 1

W (23) 4 W (20) 4 > all these OL Bh ae

W (27) | 3 N (543) 3 matings 2) oy MI GiaeO aaa

| W(14) | 14SB WwW (30)| 4 | il) RxR!| 1 1 ee

if (24) 2 5 W (30) 4 Te eae 4 1 3

| w (26) | 11 AEN a(S aaa 2 ie Bo 8 ee

W (a) 1 2 |W (8) 14SB LJ Pe 30) 2 strains.

* In both these birds the amount of webbing is just on the line fixed by

Mr. Staples Browne.

Tf we look closely into the foregoing table, we may note that

whenever the apparent Normals are mated together we get some

webs. This has been done in experiments 3, 4, and 7, the total

result, being 12 normals to 6 webs. Were these normals hetero-

zygotes we should have expected 12 Normals and 4 Webs.

WEBFOOT CHARACTER IN PIGEONS.

Text-fig. 4.

WEBBED PAIRS rrom STAPLES BROWN, EX.44. P.Z.8.1905.

W Wo W Ww WwW

No WSN N NG No Wo Wo

NNNNW/]|N NNNNW NoN NN NNN NN W

WE N N N W

Genealogical Table showing matings and results in mixed strains.

7 ss,

EXALZ. ExPJ1.

| AND MANY. OTHERS

3 ALL WEBBED.

WEBBED DRAGOON SMALLEY 69

N Wo WW NN WW

EXP AS:

WY. SW

Proc, Zoou, Soc,—1911, No, II.

18 ON THE WEBFOOT CHARACTER IN PIGEONS.

If we take the Normal-Web matings, as in experiments 2, 6,

9, 10, and 13, we find the total results to be 11 Normals to

10 Webs. In this case were the Normals heterozygotes we

should expect equality, which is practically the result attamed.

Lastly, when we mate Webs to Webs, as has been done in

Exps. 1, 5, 8, 11, 12, and 14, we should expect Webs only, and

this result, was only attained in one case (Exp. 8), where only two

birds both webbed were reared. In the other five cases together

15 birds were reared, 12 Normal and 3 Webbed. The proportions

are almost exact for a DRxDR mating, which, of course, we

know they cannot be; but it is not without significance to note

that in four out of these five experiments the matings were a

cross between the mixed strain and Mr. Staples Browne’s strain.

These, then, are the facts, and at present it seems difficult

to reconcile them with the Mendelian theory, although they

certainly seem to show that the inheritance of this character is

to a certain extent in accordance with that theory.

According to previous and contemporary work on this character,

no individual should have had a normal foot. If, however, the

Normals that have appeared be considered as heterozygotes, then,

considering the small numbers, the results from the matings are

not greatly at variance with what we should expect.

As regards the Webs (in these experiments), when mated with

Normals they behave as recessives, but when mated with other

Webs of either the mixed or of Staples Browne’s strain they

produce Normals to Webs in a ratio closely approximating 3:1.

Such a result in the case of no less than five matings seems to

show pretty conclusively that the joining of the two strains has

produced a factor disturbing the normal course of the Mendelian

inheritance.

It must be carefully noted that each strain by itself breeds

true according to Mendel’s laws and that the disturbing factor 1s

not brought in from one side only, but is due to the intermixture

of the two strains.

Mention should perhaps be made of a suggestion by Prof. Bate-

son, to whom I wrote on the subject. This was that the webbed

foot was possibly a double character and that the web between

digits 2 and 3 had a separate inheritance from that between digits

3 and 4. Now, if we look at Exp. 1 in this light, we find that

in 2 the male is very slightly webbed (practically normal) and

that in ? he is fully webbed. In the female, on the other hand,

the webbing in 3 is very full and in 2 extremely small.

We could thus suppose the male in Exp. 1 to be | N (W)

2 WW, female 3 WW 2 N(W); this would then give in F,

2 Normals and Webs equally, ? Normals and Webs equally, so

that we might expect pure Normals (; N ? W, 3 W ? N) and pure

Webs in equal numbers—as a matter of fact, 4 were Normals and

12N3W.

This suggestion, however, becomes untenable for other reasons.

If we examine the birds I bred for Mx, Staples Browne (Exp. 14,

ON THE CRANIAL CHARACTERS OF LACERTA JACKSONI. 19

loc. cit.), together with 5 more individuals reared since, we find

that the old maleno. 19 SB was undoubtedly | WW 2 N (W) and

the female was the same ; ; consequently all birds bred gual this pair

should be webbed in Zand a quarter of them webbed 1 in 2. Asa

matter of fact, out of 10 birds, 1 is normal in 3, 3 are webbed

in ?, and one fae all four toes Pilly webbed.

It is unnecessary, therefore, to follow up the matter further, as

slight deviations appear in almost every mating.

We must, therefore, be driven back to the suggestion that

a factor composed of two parts (cryptomeres), one of which is

contained in each strain, causes the web to be suppressed or

concealed.

On this assumption none of the results attained are at variance

with the Mendelian theory. The proportions, however, do not

fit in, but as the numbers are small this does not count for much.

On the average the extent of the webbing shows a tendency to

diminish and the foot to approximate to the normal as compared

with birds of either of the pure strains, and in only one bird, bred

in Exp. 9, has the web included the hallux.

Owing 40 lack of space and pressure of other experiments, I

do not propose to pursue this inquiry further, but should anyone

wish to follow it up I shall be very pleased to place all my material

at his disposal.

My best thanks are due to Mr. Smalley, who has not only

supplied me with full and accurate particulars from his pedigree

book, but has also during the past year kept and bred many of

the birds in his lofts.

2. Notes on the little-known Lizard Lacerta jacksoni Blgr.,

with Special Reference to its Cranial Characters. By

Evwarp Draen, F.Z.8.

[Received September 15, 1910: Read November 15, 1910. |

(‘Text-figures 5-7.)

Through the kindness of Mr.G. A. Boulenger, F.R.S., V.P.Z.8.,

a series of seven specimens of Lacerta jacksont has been placed

in my hands for investigation. These specimens, recently pro-

cured by Mr. R. Kemp at Mumias, Mount Elgon District,

British East Africa, at an altitude of 7000 feet, reached the

British Museum in a bad state of preservation ; opportunity was

therefore taken to submit the cranial structure of this little-

known species to a careful examination, the results of which are

now placed on record.

Of the seven specimens six were males and one female.

The specimens from which the skulls were prepared were a

small, but quite adult male, measuring 70 mm. from snout to

vent, and the single female, measuring 68 mm, The dimensions

20 MR. EDWARD DEGEN ON THE

of the former therefore are about the same as those of the single

male specimen from which the species was originally described

and figured*, after its discovery at “ Ravine Station,” Mau

Mountains, British East Africa, at an altitude similar to that at

which the specimens now studied were obtained. :

DESCRIPTION OF THE SKULL OF THE MALE.

Dimensions.—Total length 21 mm.; width 12 mm. (as measured

between the extreme points of the zygomatic processes of the

jugular bones) ; height 6 mm.

Its width is therefore contained 1-9 times in its length, and its

height 3:3 times.

Text-fig. 5.

Skull of male.

Upper, lower, lateral and posterior aspects. 22.

GENERAL ConFriGuRATION.— Remarkably depressed, as may he

noted from its proportionate dimensions, and further seen in the

above figure representing the profile.

Facial portion. Snout short, broadish at base, terminating in

a bluntly rounded-off premaxillary.

F.R.S., Proc. Zool. Soc. 1899, pp. 96 & 97, pl. x.

GRANIAL CHARACTERS OF LACERTA JACKSONI. Zi

Crantum (fronto-parietal portion).—Squarish. The whole of

the upper surface from the anterior borders of the nasals to the

posterior margin of the parietal quite flat, and forming an almost

perfectly level plane.

Dermal incrustation. Evenly spread, profuse, but highly

porous, and covering the whole of the upper surface, including a

small portion of the nasal process of the premaxillary bone.

Lamina supraciliaris (text-fig. 6, A, p. 23) (Supra-orbital

region). Completely ossified ; 4 supraoculars, and 1 supraciliary

on each side.

Supraorbital bones (text-fig. 6, B). Small; irregularly trian-

gular and thickish, convex laterally. The internal angles termi-

nating in a hook-like spine and the posterior margins irregularly

serrated. Their lateral edges barely showing on the exterior and

almost wholly concealed under the first supraciliary scutes.

Pterygoid bones. Toothless ; moderately diverging immediately

in front of the dilated lateral processes of the basisphenoid.

Postfrontals forming a single plate.

A Retrociliary present (text-fig. 6, A).

Dermal ossification of Temporal region. Supratemporalia | and

2 partially ossified.

Dentition. Number of teeth in premaxillary 9; in each

maxillary 18.

Nasal apertures (text-fig. 6, D). Subcircular, the longitudinal

diameter the greater.

Parietal Region.—This, as in all Lacertide, is clearly defined

in contour, and forms the greatest expanse of the cranial roof.

Its anterior borders practically are coincident with the fronto-

parietal suture and the postorbital margins of the adjacent post-

frontals. Posteriorly, the parietal is produced well backwards,

and ends in a straight edge, the latter being the result cf an

increased deposit of the crusta calcarea—a feature mostly absent

in the very young, in which this portion is concavely crescentic.

This edge connects the two outward and backwardly directed

processus parietales, spanning over the parotic processes, the

processus ascendens of the supraoccipital bone, and covering the

greater part of the latter in this species, so that the pars condy-

loidea of the occipital alone projects beyond it when the skull is

viewed from above. .

The parietal processes, slightly triangular in a transverse

section at their terminal parts, are quite flat at their basal

portions and are mere continuations of the median portions of

the parietal itself. They have not, as is the case with the

majovity of the Wall-Lizards in which the facies cranialis is more

elevated, the downwardly directed ridges set at right angles with

their bases, in order to establish the contact with the underlying

supraoccipital,

No Foramen parietale. (For special remarks on this striking

feature refer to p. 29 at the end of the description cf cranial

characters.)

he MR. EDWARD DEGEN ON THE

The Postfrontals, which are joined laterally to the single parietal

bone by means of ordinary sutures, and which extend the whole

length of the parietal as far backwards as the supratemporal

bones, complete the cranial roof. They form a single plate in the

present species, with every trace of their former sutures com-

pletely obliterated through fusion of the two component parts

of which they consist in a great number of Lizards. This is a

condition found to exist also in Lacerta simonyi, L. atlantica,

L. ocellata, L. viridis, L. gailoti, L. dugesii, and L. levis (in

the latter species the squamosal also fuses with the postfrontal

plate). In Lacerta jacksoni these bones are on the same hori-

zontal plane with the parietal to their outermost margins, which

in so many of the Lizards of the “muralis” group are strongly

convex and appreciably bent downward towards the temporal

foramen.

A similar feature, which imparts such a quadrangular

aspect to this portion of the skull, is observable in the case

of Lacerta simonyi*, L. atlantica, L. ocellata, L. viridis, L. galloti,

L. dugesii, L. levis, Algiroides nigropunctatus; and among

L. muralis forms in the varieties nigriventris and serpa.

ally paired (in very young specimens) it is

single in the skull before me as well as in the majority of the

members of this genus when adult. Its interorbital width, at

the narrowest part of its anterior portion, measures exactly one-

half of its posterior and widest portion, which is to be found in a

line drawn between supraocularia 4, the latter coinciding approxi-

mately with the fronto-parietal suture. In proportion, therefore,

this interorbital width is greater than in most of the mwralis-like

forms, except for the skull of a male specimen of the typical form

from Vienna, which is also greatly depressed. In the length of

the skull sans dimension is contained six times; whereas in the

majority of skulls belonging to the mwralis group it varies

between seven and eight times. It further equals the width of

supraocularia 2 and 3; also that of the space between the inner

dentary ridges (laminz horizontales) of the maxillary bones

(measured across the apertura narium interna), and is of the

same length exactly as the columella cranii (epipterygoid, Parker).

Equally distinguished in respect to this broad interorbital

diameter are—apart, again, from all the previously enumerated

Atlantic Island forms and others named—the skulls of Z. agilis

and levis, and of the more typical muralis forms those of

L. chlorogaster, L. saxicola, and L. derjuginit. Further, in

Algiroides nigropunctatus this region is also of a broader type ;

but according to Siebenrock¢ it differs in the persistence of the

SCE “Das Skelet der Lacerta simonyi Steind. und der Lacertenfamilie iiber-

haupt,” von Friedrich Siebenrock, Sitzungsb. Kais. Akad. der Wissenschaften in

Wien, vol. ae parti., April 1894, Taf. 11. fies. 8 & 11.

+ fr. L . von Méhely, “ Materialien zu einer Systematik und Phylogenie der

Muralis-alnlichen Lacerten,” Ann. Mus. Nat. Hungarici, Band vit. 1909, tab. xix.

fic. 9 & tab. xxi. figs. 3 & 6.

tT esp. 28.

CRANIAL CHARACTERS OF LACERTA JACKSONI. 23

two original parts composing the single frontal in the adult of

this genus also *.

Text-fig. 6.

A. Lamina supraciliaris of Lacerta gacksoni, 6. Right side.

B. Supraorbital bone of ZL. jacksoni, g. Right side, ventral aspect.

C. Supraorbital bone of ZL. jacksoni, 2. Left side, ventral aspect.

D-I. Showing configuration of nasal process of preemaxillary and outer nasal

apertures of —(D) L. jacksoni, 6; (KE) L. jacksoni, 2 ; (F) L. muralis, f. typica,

d@, from Vienna; (G) L. muralis, f. typica, 6, from Turin; (H) L. vivipara, fg ;

(1) ZL. agilis, $. X 65.

prf. prefrontale; so., seuta supraocularia (1-4); se., seutum supracihare ;

s.r., scutuim retrociliare; s.o7., supraorbital.

Premacillary.—lts backwardly directed nasal process (text-

fig. 6, D) is rather short and exceptionally broad at_ its base,

where it is widely expanded on both sides, close behind the

* The skull of the male specimen of this species in the British Museum Collections

does not bear out this statement, the said suture being wholly obliterated by

incrustation,

24 MR. EDWARD DEGEN ON THE

laterally situated nerve-canals which at this point perforate the

bone. Other examples of a very broad nasal process of the

premaxillary are to be found in Lacerta ocellata, L. viridis,

L. agilis (text-fig. 6, I); and reference to Dr. Siebenrock’s

text and figures for L. simonyi* also shows the latter to belong

distinctly to the broad type, and consequently not—as he points

out, when referring to this feature—being distinguished by

a “long and slender” type of processus nasalis the same as in

Eremias arguta, with which he likens LZ. simonyi as regards this

character.

In Lacerta dugesii I find the shape of the nasal process occupies

a position intermediate between ZL. jacksoni, the species here dealt

with, and those possessing a nasal process of distinctly slender

and elongate shape. Most of the true mwralis forms belong to

the latter (text-fig. 6, F & G). Lacerta echinata, L. vivipara

(text-fig. 6, H), and Z. devis must also be considered intermediate

forms in this respect. Allowance for individual variation, how-

ever, must be made, as the examination of three skulls of the

latter species, all males, clearly demonstrates.

The very low angle at which this nasal process rises at its

issue from the maxillary portion of the bone is merely due to

the strongly depressed type of the skull itself, as, in fact, in

any of the typically platycephalous forms, e. g. ZL. mosorensis or

L. oxycephala, and differing from the more or less convex types

of skulls in which this process is more highly arched.

Nasal bones.—Owing to the profuse incrustation of the upper

surface of the skull, their contours in this specimen are very

ill-defined. All that can be said on this head is that they

are oblong, of a scaly nature, and that they widely diverge

behind to receive between them the anterior and pointed pro-

cessus nasalis of the frontal bone. Their length approximately

equals their combined width at the widest part between the

prefrontal bones.

The Prefrontals also appear very irregular in their upper out-

lines and in their relations to the maxillaries, the frontal, and the

nasals, The spinous posterior process, which forms the anterior

border of the margo supraorbitalis, seemingly does not reach much

farther backward here than the suture between the first and the

second supraorbitals.

The Zacrymal bones are clearly defined, exceedingly thin,

narrow, and blade-like in shape. In their situation at the

anterior angle of the orbital cavity, they form, together with the

incisura lacrymalis of the prefrontals, a well-developed foramen

lacrymale.

Jugulars.—Their zygomatic processes are represented by a

rather short, but acutely pointed spine. Their temporal processes

are somewhat straighter than is the case in Lizards having

deeper skulls, in all of which they usually are of a more sigmoid

form,

oe 156.65 ie BPA, Weis Tub, sly, Eh

CRANIAL CHARACTERS OF LACERTA JACKSONT, 25

OccIPIfAL SEGMENT.—Supraoccipitai. ts epiotic portion is

neatly horizontal and only slightly inclines towards the posterior

free edge, which forms the upper margin of the foramen magnum.

The processus ascendens (processus spinosus, Clason) rises rather

abruptly and almost vertically from the anterior portion of the

supraoccipital, and constitutes the exclusive support of the

parietal roof.

The processus ascendens is not flanked on either side by any

apophysial eminence of the supraoccipital, which so frequently

acquires the same height as does the spinous process itself, in

order to increase the connecting-ar ea with the pemenal, such as is

the case in Lacerta oxycephala, L. saaicola, L . defilippi*, L. chloro-

gaster (boettgert Méh.), and L. mosorensis, among some of the flat-

headed species; or as in Lacerta muralis forma typica, the

varieties tiliguerta, campestris, niyriventris, &c., as well as Lacerta

vivipara, among the more highly and convexly-roofed forms

where it also occurs.

A similar pillar arrangement as pointed out for Lacerta jack-

sont is present also in ZL. stemonyi, where it is carried out to

even greater perfection owing to the inereased elevation of the

parietal roof +; in L. galloti, L. viridis, L. dugesii, and L. echinata,

proportionate, of course, to their respective dimensions of height.

This arrangement is of rare occurrence in a strongly depressed,

practically platycephalous Lizard, and equally so in the case of

the more pronounced pyramidocephalous forms, amongst which

I find it occurs (in two specimens) in the varieties serpa and

lilfordi only.

The Basioccipital £ and Basisphenoid § in no essential deviate

in their configuration from the outlines usual in the genus for

this region.

The Parasphenoid is feebly ossified at its base, whereas its

centre remains cartilaginous.

The Orbitosphenoid bones have a broadish appearance on

account of their reduced size, the latter being due to the depressed

membranous portion of the cranium.

The Squamosals, usually slender and delicately shaped, which

in so many of the smaller forms of Lacerta are only loosely con-

nected by ligamentous tissue with the outer postfrontals, are here

strongly developed and anteriorly quite fused to the latter.

Their posterior ends, which articulate with the quadrates, are

broadly expanded and, together with the parietal processes and

the much reduced supratemporals which wedge themselves

* Cfr. Méhely, “ Materialien etc.,” Ann. Mus. Nat. Hung., Bd. vii. 1909, tab. xvi.

Hh 6, tab. xix. figs. 3 & 10, tab. xxii. fig. 6.

+ Ofr. Siebenrock, /. c. Taf. ii. fig. 9.

t Posteriorly and slightly in front of the foramen magnum there is a small

aber cular process in the male specimen, which is absent in the female.

§ The foramen by which a branch of the jugular vein leaves the skull, and which

Siebenrock (/.c. p.11) regards as the result of synostosis of a special process with

the anterior angle above it in the case of the Canary Island Lizards, but as absent

in the European forms as well as in ZL. duyesii, is represented in L. jachsoni by

ligament only. :

26 MR. EDWARD DEGEN ON THE

between these two latero-posterior angles of the skull, unite

with the processus parotici in such a way that the foramen supra-

temporale, conspicuous for its relative large size even in the

adults of the pyramidocephalous forms like Lacerta muralis

varieties jfiumana, serpa, campestris, pityusensis, and others,

is here nearly closed up as much, for instance, as in L. levis.

The Presphenoid, which in some Lacerte is a slender ossifica-

tion of the ethmoidal region in the membranous septum orbitale,

is here reduced to a small cartilaginous band in which calcifica-

tions in the shape of granules are imbedded.

The Columella cranii, for a skull of the inferior dimensions

of the present one, are exceptionally thick and strongly bent

laterally—particularly so at their parietal ends, where they are

m close contact with the ala sphenoidea. Thus they differ

somewhat from the more delicate, either linear or slightly sig-

moid thin structures frequently met with in the majority of other

Wall-Lizards ; but this condition is the rule in the very young,

as already pointed out by Siebenrock *.

The Quadrata and Ossa transversa both answer to the general

form of description for all these Lizards.

The Vomer plates are closely approximated to each other

medially along their’ entire length, but intimately united only

at their anterior ends. The lacuna pterygo-vomerina does not

extend beyond the most posterior portion of the palate-bones,

which latter also unite with each other along their greater length,

producing the crista vomerina which is continued on to the

palatines.

Manvi1BLe.—This consists of five elements only,a condition stated

by Leydig to be typical of the Lacertw, and further confirmed also

by Siebenrock fT in the fully adult Zacerta atlantica, L. muralis

var. melissellensis, L. cerulea, L. oxycephala, and L. mosorensis,

a condition brought about by the fusion of the supra-angulare

with the articulare; whereas in L. simonyi, ocellata, galloti,

and viridis t, according to Siebenrock, these two bones remain

separated throughout life.

The number of teeth in each ramus is 23.

DESCRIPTION OF THE SKULL OF THE FEMALE.

Dimenstons.—-Length 15 mm.; width 9 mm.; height 4 mm.

Its width is therefore contained 1-6 times in its length, and its

height 3°3 times.

GENERAL CoNFIGURATION.—Strongly depressed in the same pro-

portions as in the male.

In texture the bones are rather delicate, such as is character-

istically the case in all platycephalous species of Lizards so

* Tc. p. 52. p ip Jey @ ite BS:

'£ In the skull of a specimen of this species trom Bozen, Tyrol, these two hones,

though fused slightly, show their original longitudinal sutures in their anterior

portions.

CRANIAL CHARACTERS OF LACERTA JACKSONI. 27

typically represented by Lacerta mosorensis, oxycephala, ke.

Except for the bones composing the occipital segment, and the

usually more massive premaxillary and maxillaries, all the others

assume a more or less foliated form.

Facial portion. 'The snout appears much shorter and broader

than in the larger male, but is slightly more pointed in front.

Text-fig. 7.

Skull of female.

Upper, lower, lateral and posterior aspects. X 3.

Cranium (Hrontc-parietal region).—Slightly broader than long

in proportion, owing to the incomplete development of its posterior

margin, which is crescentic in shape as, in all young examples.

Its median edge therefore barely reaches the anterior margin of

the supraoccipital. The upper surface, quite horizontal in its main

configuration, is longitudinally slightly undulated by alternately

being concave in its fronto-nasal and fronto-parietal parts re-

spectively, and convex in the intervening frontal and parietal

areas.

Dermal incrustation. Very sparse and transparent. The sulci

of the blood-vessels are sharply defined for their former rami-

fications.

Lamina supraciliaris. Completely ossified also, as in the male,

and not bearing a trace of a membranous fontanelle.

28 MR. EDWARD DEGEN ON THE

Supraorbitals (text-fig. 6, C, p. 23). More regularly triangular

than in the male, alsoslightly more massive. Their thickened and

strongly convex lateral margins, scarcely exposed in the male, are

here so to a greater extent on both sides (ascertained before

accidental displacement of their dermal plates on one side in the

later manipulation of the skull), having been covered partially

only by the supraciliary.

Pterygoids. Diverging slightly more than in the male speci-

men. ‘Teeth absent.

Postfrontals. Forming single -plates also. Faint traces of an

earlier existing suture, as recorded to exist in the very young

stages of Lacerta dugesit, ocelluta, and muralis by Siebenrock*,

are discernible anteriorly only. In their posterior portions

(postorbital margins) their fusion 1s complete.

A minutely developed Retrociliary present.

Dermal ossifications of Temporal region. I have been unable

to find any.

Dentition. Number of teeth in premaxillary 9, in each maxillary

15-16.

Nasal apertures (text-fig. 6, KE). These ave slightly narrower

than in the male, and therefore longer in proportion.

Parietal Region.—Except for its morphogenetically inferior

development of the posterior margin, which is a common feature

of all the young, this does not deviate in any essential from the

usual conditions. Its relation to the supraoccipital alone requires

some explanation (see p. 29), which latter bone is, owing to the

development stage, wholly exposed and not yet bridged over as

it is in the male.

The Foramen parietale is absent, as in the male.

Frontal—The median suture is discernible in its greater

length, which is quite in accordance with the remarks aleady

made on this subject. The width of the frontal at its narrowest

point (between the margines orbitales) is equal to that of sapra-

oculary 3. The frontal is actually and relatively narrower than

that of the male, its width being slightly less than half the

length, and a seventh of the total length of the skull, whilst in

the male the similar measurement shows that the frontal is

relatively larger. ‘The width of the female frontal is three-

fourths of the length of the columella cranii, and in this measure-

ment, as well as when it is compared with the width between

the internal Jamine horizontales (taken diagonally across the

choane), the relative and actual inferiority of size as compared

with the male is shown.

Premaxillary (text-fig. 6, E)—The nasal process of this bone

is on the whole longer and narrower than in the male. It also

terminates posteriorly in a much more finely drawn-out point.

It is quite devoid of any calcareous incrustation. Anteriorly it

rises from the maxillary portion of the bone at a similar acute

“OL cupe ios

CRANIAL CHARACTERS OF LACERTA JACKSONI. 29

angle to its horizontal plane as in the male, but is less expanded

at its base, immediately behind the constriction between the

nerve-holes.

The Nasal bones, which, owing to extensive incrustation, in

the male specimen were described as ill-defined, are here quite

distinct in their entirety. Anteriorly their proximal processes,

between which the nasal process of the preemaxillary is wedged,

extend to the middle of the latter. Their lateral margins, shorter

than those forming the median suture and moreover less decidedly

linear, are parallel to each other. ‘Their posterior edges, scale-

like in shape, and imbricating in a similar manner on the frontals,

are characteristic in having from 2 to 3 pointed lobes, of which

the median ones are the longest, so as to become a fork into the

prongs of which the median process of the frontal penetrates.

The Prefrontals also are more clearly defined in the female.

They show their posterior frontal processes, which constitute the

margo orbitalis anteriorly, to be spurs which extend nearly as far

back as the suture between supraocularies 2 and 3.

The Jugulars do not differ from those of the male.

OcciprvAL SEGMENT.—The Supraoccipital is united along its

entire upper margin with the posterior edge of the parietal bone.

The processus ascendens, so tolerably well developed and forming

the sole support of the parietal in the male, is quite rudimentary

here and reduced to atubercle only. It is lower even than are the

apophysial elevations of the epiotic portions of the bone, and,

together with the epiphysial cartilaginous spine and a small

portion of the membranous cranium, is laid bare and quite un-

protected by the parietal. The sutures between the supraoccipital

and the pleuroccipitals are perfectly distinct. The foramina

retrofrontalia, in consequence of this extended transverse con-

nection between the parietal and the supraoccipital, are more

markedly reduced than in the male.

The Basioccipital and Basisphenoid are not fused as in the male,

the suture between them showing distinctly (text-fig. 7, p. 27).

The Squamosals are flatter and less curved downwards than in

the male.

The Vomer plates are more loosely united than in the male ; but

the Palatines, though more distinctly united, have their sagittal

ridges imperfectly developed as compared with those of the

male. The swlct palatine are also shallower in this skull.

ManpisBLe.—This consists of the six elements, of which each

ramus is composed in all immature Lizards as pointed out

by Siebenrock, the supra-angulare and articulare being not yet

united by fusion.

On the Absence of the Foramen parietale.

After first ascribing the absence of the orifice for the pineal

eye in the skull of the male to purely pathogenic causes, or to

30 MR. EDWARD DEGEN ON THE

a possibly individual aberration, both of these assumptions were

dispelled on the discovery of this peculiarity in the female aiso.

An examination of the remaining five specimens preserved

in alcohol, as well as of the type specimen referred to (antea,

p- 20), and of a second female more recently described * (all

in the Collection of the British Museum), readily confirmed its

specific importance, since not one of them bore the least trace of

the presence of this morphological feature.

Besides the simultaneous observation of its absence in Zono-

saurus madagascariensis, belonging to the neighbouring family

Gerrhosauride, a more methodical search among the rich

material of the British Museum Collections revealed the following

genera and species belonging to the family of Lacertide as being

devoid also of the foramen parietale :—

Poromera fordii. Benito River, Spanish Guinea.

Lacerta echinata. Kyibi River, Cameroon.

Algiroides africanus. Uganda,

Nueras delalandi. Natal.

Latastia hardeggert. Somaliland.

- NEWMaANNE. Arabia.

- degen. Somaliland.

a spinalis. .

» prillipsir. A

Gastropholis vittata. Zanzibar.

Holaspis guentheri. Benito R., Spanish Guinea.

As the above list shows, cases of absence of the parietal

foramen occur in several monotypic genera, also in the specifically

numerous genus Latastia in no less than five species out of ten.

Yet it is rather remarkable to find such a character restricted to

two species only in a large genus such as Lacerta.

But more remarkable, perhaps, is the fact that all the forms

thus deprived of the foramen belong to the African Continent,

and that in the not strictly African genera it is restricted to

African representatives.

Some of the remaining genera of this family, even of purely

African habitat, do not contribute any specific forms which exhibit

this negative character. It is curious to note, moreover, that

among the upwards of thirty known species of the genus Hremias,

about twenty of which are found in Africa, no exception to the

presence of the parietal foramen could be found.

The sum total of cranial characters, as shown principally in the

immature skull of the female, points to Lacerta jacksoni being

* G. A. Boulenger, “ Ruwenzori Expedition Reports,” Trans. Zool. Soc. vol. xix.

part i1., Dec. 1909, p. 242.

CRANIAL CHARACTERS OF LACERTA JACKSONI. 5)

closely allied to Z. muralis, forma typica. It agrees with the

latter as well as with Z. mosorensis in the strongly depressed and

more broadly expanded skull-structure in general. Also in the

wider interorbital region, likewise a marked feature in Lacerta

vivipara. Also in the mode in which the connection between the

posterior portion of the parietal and that of the supraoccipital

presents itself; in the absence of teeth on the pterygoid bones,

and in the degree of divergence of these latter, as well as in the

configuration of the basioccipital and the basisphenoid. Further,

in the shortened muzzle with broadened base, and in the

anteriorly broadened nasal process of the premaxillary bone

(text-fig. 6, D), well indicated in the typical form of LZ. muralis

from Turin (text-fig. 6,G), but assuming its widest proportions in

Lacerta agilis (text-fig. 6, 1).

The total absence of supraocular fontanelles, notwithstanding

the typically platycephalous structure, together with the onto-

genetically early complete fusion to one composite plate of the

postfrontals, would justify our referring this type of skull to the

group of Lizards to which the term of Neolacertie * has recently

been given,

However, considering the views held and recently enunciated

by writers on the subject of the phylogenetic relation of some

of these characters, it is rather surprising to find undivided

postfrontals also in species belonging to entirely different genera,

such as Hremias, as Prof. v. Méhely ¢ regards that genus as one

of the prototypes of archeolacertic forms of this family because

of the characters of its lepidosis.

I might add that in Acanthodactylus boskianus{, a lizard to

which Siebenrock$ ascribes a divided condition of the post-

frontals, I find them to be completely fused into one plate in the

skull of a female in the British Museum Collections.

A perfectly obliterated suture between the two postfrontals

I also find in a female specimen of Scapteirw knowii, another

decidedly platycephalous species.

Thus, beyond some more or less doubtful points in regard to a

satisfactory solution concerning the question of phylogeny, skulls

of females do not seem to afford trustworthy indications from

which to draw conclusions as to affinities. It is to that of

the male, so far as an examination of the present limited

material is concerned, that one must turn for better results.

As pointed out in the detailed part of the description, the

tabular surface (inclusive of the postfrontals) is a very conspicuous

feature of this male skull when taken in conjunction with the

quadrangular and posteriorly well projecting parietal region. A

near approach in this respect is to be found in a male specimen

of Lacerta muralis of the variety lilfordi, a form restricted to

* L.v. Méhely, Materialien, ic:

£ Sex not stated by Siebenrock.

§ Cf. “Das Skelet der Lacerta simonyi,” Sitzb. Akad. Wiss. Wien, 1894, p. 37.

32 MR. EDWARD DEGEN ON THE

the Balearic Islands. Though unquestionably more pyramido-

cephalous in regard to its superstructure, it nevertheless holds a

position somewhat intermediate between Z. jacksoni and some

of the more pronounced pyramidocephalous forms in this group

in regard to this particular feature.

The variety lilfordi also shows the same exclusive pillar arrange-

ment of the processus ascendens for sole support of the parietal

roof, which is so prominent a feature also in all the larger forms

of Lizards from the Atlantic Islands such as LZ. simonyi, galloti,

atlantica, and also L. dugesit* (except in regard to a modification

in the basal portions of the parietal process), all differing by having

pterygoid teeth, and all of which, except the latter, have strongly

ossified temporal regions. These dermal ossifications in Lacerta

jacksont, as may have been seen, are present in a rudimentary

state in two of the supratemporal scutes. Specimens of Lacerta

viridis, agilis, and levis may be quoted as further examples of

characteristic forms in which the postfrontals are single plates,

with perfectly analogous conditions in the relations between

parietal and supraoccipital.

Also Lacerta echinata, from West Africa (except for its more

elongate skull), in which the processus ascendens, typically pyra-

midal, does not enter into direct contact with the parietal, but is

actually separated from it by confluence of the foramine retro-

frontalia.

But it is the shape of the nasal process of the premaxillary

perhaps, as previously remarked on in the male, which may

also provide a likely indication of closer affinity with some of

the Lizards of the Canary Islands and the European species of

Lacerta, viz. ocellata, viridis, and agilis—a character in which the

Balearic variety lilfordi does not share, being distinguished by a

narrower and very slender nasal process.

So far as the topography of the skull reveals phylogenetic

affinity, it would appear that Lacerta jacksoni came from a

Palearctic stock, but also exhibits special relationships between

the West African and Atlantic Island forms.

Remarks on Characters of Lepidosis in Lacerta jacksoni.

Dimensions.—In regard to size, five specimens, all of them

males, are larger than the type of the species; the two biggest

measuring 86 mm. from snout to vent, and consequently of

about the same length as is the largest of the males of Dr.

Peracca’s series (consisting of seven specimens also), the length

of which is given as 85°5 mm.

The male specimen, from which the skull was prepared for the

* Siebenrock (vide l. c. p. 36) includes this species amongst those having divided

postfrontals. ‘The specimens at my disposal, from the British Museum Collections,

show them distinctly to be single plates, with no indication of former sutures.

CRANIAL CHARACTERS OF LACERTA JACKSONI,. 3o

foregoing description, measured 70 min. ; equal to the length of

the type specimen. Jt is therefore only slightly longer than the

skull of the female which was sacrificed for the same purpose, the

length of the latter from snout to vent being 67 mm. ‘Though

differing by 5 mm. only, the sexual discrepancy in the size of the

heads is a striking one: 19 mm. for the male against 14 mm. for

the female. ;

Analogous specimens for sfze in Dr. Peracca’s series show

corresponding difterences, viz. 20 mm. for the head of a male

and 15°5 mim. for a female.

Ventral Plates —The number of the ventral plate series varies

in the Mount Elgon specimens, three of these having 6 longi-

tudinal rows only ‘and four of them having 8, which is the same

as in the type of the species.

For the Turin Museum specimens their number is uniformly

stated to be 6; but to judge from a footnote im connection with

this point, the author was fully cognisant of the presence of

supplementary series* for some of his specimens, but hesitated to

accord them serial rank on account of the inferior size of the scutes

which form the outermost rows.

In several of the Mount Elgon specimens these latter are

sufficiently well developed to be considered as a series of ventrals.

Transversely, the minimum of the plates in the Elgon speci-

mens is 23 in an adult male, and the maximum 26 respectively

for the male and the female from which the skulls have been

prepared.

For the Turin Museum series this minimum of 23 occurs in

two males, one quite young and the other larger, whereas for

the largest and fully adult the number is 24, ‘but the highest

figures oe 27 and 28 are reached by two femeles ot the same lot ;

the first number being identical with that of the Ruwenzori female

in the British Tinea Collections, whereas the female specimen

from Mount Elgon has only 24 rows of these scutes.

It must be borne in mind that female lizards, having the body

more elongate in proportion than the males, usually possess a

higher number of transverse series of ventral plates.

Seales across the middle of the Body.—'these range between

38 and 43, giving an average of about 40, the same as recorded

for the type specimen, provided we except the male from which

the skull was prepared and which showed the abnormally high

number of 49. The Ruwenzori female has 37, a number

close to the average of 36 as represented in the Turin Museum

series, Where they vary between 34 and 38, the latter figure

being that also of Lacerta vauereselli, as well as constituting

the minimum of the British Museum Elgon series of Lacerta

jackson.

Supraciliary granules.—These vary in Oe specimens, both

individually and for the two sides, between 3-3 and 6-5. They

* Ofr. Dott. M.G, Peracea. Tl Ruwenzori, Relazioni Scientifiche, vol. i. Goologia-

Botanica: Rettili ed Amfibii, Lacertide, pp, 166 & 167.

Proc. Zoon, Soc, —1911. No. ITY, 3

34 MR. EDWARD DEGEN ON THE

are confined principally to the posterior portion of the supraocular

region. In the Ruwenzori female they are reduced to 3 on the left

and to 2 on the right side. Dr. Peracca describes these as “ very

irregular and asymmetrical for the two sides, except in a female in

which the series is a complete one on the teft, but totally absent

on the right side.” According to Tornier, who in his description

omits to quote their number, the supraciliary series of granules is

incomplete also in Lacerta vauereselli; but, as he says, confined to

the posterior portion, agreeing therefore in Has point also with

L. jackson.

rular Scales.—The number of scales between the third chin-

plates and the collar-plates varies between 24 and 26 in the Elgon

specimens except in one, in which there are 28. This is slightly

in excess of the Duke of the Abruzzi’s Expedition specimens, in

which they are stated to be 22. In the type of the species

their number is 25.

Femoral Pores. —Vheir number is fairly constant, with variations

between 16 and 18in the Elgon series. In one of the two females

of the Turin Museum series there is a reduction to 15-14, otherwise

17-18 being the normal number. No case of supernumerary pores

occurs in any of the British Museum specimens similar to the one

female quoted in this respect by Dr. Peracca.

Upper Labials (anterior to the subocular).—Except for the head

of the male, made into a skull, which on its left side showed these

to be 5, their normal number throughout the remainder is 4.

A male of Dr. Peracca’s series shows a similar variation.

Postocular and Temporal Region.—In one of the specimens the

last postocular scute is in contact with the first supratemporal,

similar to what is found in Lacerta mosorensis as figured by

Méhely *. In all others it is the upper postocular and the parietal

scutes which meet, as in Lacerta horvathi Méh. or L. muralis

typica T

The 3 to 4 supratemporals are exceedingly small and narrow,

the first and longest being partially supported proximally by

the lateral margin of its underlying postfrontal bone, and

therefore visible practically in its greatest extent when the head

is viewed from above.

The number of scutes for the temporal region varies greatly.

Those, for instance, forming the second row between the large sub-

ocular and the first supratemporal are: 3 for the left, and 5

for the right side in the male, the skull of which served for

description.

Similar variation is to be found amongst the scutes in regard

to their sculpture, which shows every gradation, from the per-

fectly smooth through the granular stage, as pointed out in Mr.

Boulenger’s description of the typical specimen ¢, to the faintly

keeled or tubercular scutes to be met with in some of the Elgon

* Ofr. Ann. Mus. Nat. Hung. ii. 1904, p. 366, fig. 2, A, “ Hine neue Lacerta aus

Uncen”

+ Ib. figs 2, B& C:-

t CfG. A, Boulenger, Proc. Zool. Soc. 1899, p. 96.

De ho

CRANIAL CHARACLERS OF LACERTA JACKSONT, 35

specimens, a condition said by Tornier to exist also in his Lacerta

vaueresella,

The suture between the first supraciliary and the second is

slightly oblique.

The frontal shield, which also varies slightly in length, and

which is especially short in some specimens, is on the w hole shorter

and broader than would appear from the figure on the plate

accompanying the description of the type of the species. It

was particularly broad in the male specimen which supplied the

skull for description.

Colour-pattern and markings are quite uniform for the Elgon

specimens, and identical with the type of the species. In their

nearly fresh state of preservation their ventral sides were of a

vivid yellow, which, since their preservation in alcohol, has changed

into the whitish blue-grey described by Tornier in his Lacerta

vauereselli, or the pale blue ascribed to the Ruwenzori female.

Remarks on some of the supposed Specific Characters of

Lacerta vauereselli Zornier.

On examination of specimens in a comparatively large series,

as may have been seen from the foregoing, it appears that some

of them are subject to individual var iations.

This is particularly noticeable in the scaling of the temporal

region, which was made one of the principal features by which

to distinguish Lacerta vauereselli from L. jacksont.

In the majority of specimens of the latter, these scales cor-

respond with Dr. Tornier’s description of them *, and a special

note on this subject made on the male specimen—previous to re-

sorting to the excision of the skull—was to the effect that these

scales were considerably larger than those of the upper dorsals,

being oblong, irregular in size, hexagonal to pentagonal in

shape, feebly though distinctly keeled, and decidedly double the

size of those situated immediately behind the auditory opening—

all of these being the characteristics ascribed by Dr. Tornier to

his Lacerta vauereseili, and which are also referred to by Mr.

Boulenger f at the end of his description of the Ruwenzori female.

iveeor ding to Dr. Tornier the collar is serrated. This, again, 1S

a point not always easy of recognition, for some specimens show

various degrees of unevenness in this respect.

The same remark applies to the number of plates forming the

collar-edge, which are stated by Tornier to be 11. In the type of

the species they are stated to be 10, which is the number also in one

of the Elgon specimens; in the remainder of which, however,

they range downward to from 9 to7 only, thus showing con-

siderable fluctuation. Peracea omits to quote their number.

* Ofr. Zool. Anzeiger, Bd. xxv., Oct. 1902, p. 702.

+ “ Ruwenzori Exped. Reports,” Trans. Zool. Soc. vol. xix. part iii., Dec. 1909,

p. 949,

lon

ON THE CRANTAL CHARACTERS OF LACERTA JACKSONI,

Nor can I find any differences in the style of scaling of the

upper dorsals between Dr. Tornier’s diagnosis and that given

for the type of the species by Mr. Boulenger, which scales, according

to the latter, and further verified by myself, are stated to be

‘“‘rhomboidal, keeled, juxtaposed, or subimbricate,” every one of

these conditions occurring in the Elgon specimens as well as in

Lacerta vauereselli.

The number of scales across the body, viz. 38 in the latter, also

is in accord with the mean of the Elgon series of specimens; and,

as regards the femoral pores, this has been finally disposed of,

both by what has been made known (andtea, p. 34), as well as by

a remark made by Mr. Boulenger in connection with this point

in his description of the Ruwenzori female.

As already pointed out in the diagnosis of the skull of Lacerta

jacksoni, there are no teeth on the pterygoids any more than

in LZ. vauereselli. No difference appears to exist in regard to

the supratemporals as described for this species by Dr. Tornier

from their condition in the Elgon specimens, in all of which a small

tympanic scute is plainly v isible.

Neither is a distinct gular fold recognizable in all specimens, at

least in the stricter sense, as it is orem apparently absent in im-

mature individuals, such as seems to be the case with Dr. Tornier’s

type, judging by the dimensions given by him—an assumption

which is further corroborated by the fact that its sex has not been

mentioned.

The length of the posterior extremity also varies greatly

individually—in some Elgon examples it scarcely attains the

axillary pit, whilst in others it reaches well bey ond the collar-

edge.

The geographical range of this species, so far as the increased,

but at present still scanty, material on record permits one of

judging (17 specimens, including Dr. Tornier’s L. vaueresella),

appears to be a strictly equatorial one.

Its habitat, moreover, seems to be generally restricted to the

mountainous region of that part of the Central African plateau,

or immediately west of it, which culminates in its highest peaks,

an area generally acquiring considerable altitude.

Its ultraplatycephalous shape, concurrently with its sombre-

coloured garb, at once seems to denote a form occupying high

ground, differing from those of the plains, with generally more

massive skulls Aad more vividly contrasting colour - patterns,

in the same manner as, for instance, eee, MLOSOTERSIS differs

from Lacerta muralis var : 2 Sanne is or serpa, and Lacerta muralis

var, monticola from the var. bocagit.

2 3 3 = = NC antl dS nade ~ + 2 5 ~ ., “a :

iS eee ee aaa

eT Quo day) }9 [Sp Ussdy Pp

ON THE PELOPONNESIAN LIZARD. 37

3. On the Peloponnesian Lizard (Lacerta peloponnesiaca

Bibr.). By G. A. BouLencsr, F.R.S., V.P.Z.S.

{Received October 4, 1910: Read November 15, 1910.]

(Plate I.* and Text-figure 8.)

Examples of a Lizard from Greece, which had been confounded

with Lacerta muralis or with L. tawricat until shown by Dr.

J. de Bedriaga ¢ to be fully entitled to specific rank under the

name of L. peloponnesiaca Bibr. §, have been exhibited for the

first time this summer in the Society’s Reptile House. As regards

coloration, the species is one of the most beautiful, and as the

only ficures of this Lizard we possess are quite inaccurate, or taken

from specimens preserved in spirit, I requested Mr. J. Green to

make coloured pictures from the specimens living in the Gardens ;

at the same time I seized this opportunity of drawing up a new

description of this little-known Lizard, with a view to fixing its

correct position in the genus Lacerta.

In addition to the live specimens I have had the following small

series at my disposal, all eee no. 3 being preserved in the

Natural History Museum :—

1-2. ¢2 Morea. Dr. J. de Bedriaga.

Onli. ae (Lataste Collection).

4-8. @Q her. LL. Stymphalos. Norman Douglass, Esq.

S10, ge Olympia. Hr. Lorenz Miller.

Tile gs Kalamata.

The skull of a male from L. Stymphalos has been prepared, and

agrees in all essential respects with that of Z. tawrica.

Form and Proportions.

As may be seen on comparing the following table of measurements

with that given for Z. tawrica||, the general proportions are the

same as in that species, except for the rather longer limbs. In

the male the hind hmb reaches the collar or a little beyond, in the

female it reaches the axil or the shoulder. The head is less

flattened than in any of the forms of Z. mwralis, closely resembling

in shape that of the typical form of ZL. viridis; snout sometimes very

obtuse, sometimes very pointed, as long as the distance between

the eye and the ear-opening; neck as broad as the head, or a

little broader. The foot is always longer than the head.

For explanation of the Plate see p. 49.

For a deseription of this Lizard see P. ZS. 1907, p. 557.

Zool. Anz. 1883, p. 216, and Abh. Senck. Ges. xiv. 1886, p. 315.

In Bory de St. Vine. Expéd, Sc. Morée, ii. p. 66.

i) Pp: ZS. 1907; p.56e:

Cott 3

38 MR. G. A. BOULENGER ON

Measurements (in millimetres).

3: a

fig te er go oy Se fee aN

123 4 5. 6G. Me tS eos LO!

IDO HO) WOM iococrecsnoo SO) 7 70 7 Th WD Wy Wl WD Oe

Pr Ay Oe, bb ego «Ge 28) RS) a) Bib Br 28 26 25 25

TSA Oe WEN 5 5 Goons os oo BL FD 19 I I BW 1G 1G 16 Le

AWGN @rP INGEN .y coseno nue ce2ane Wey Ts) es eye HO) NO) 9

IDyeyotelh Gre MEAL ceo oomsbaccesconen = IL IK) ul NG 9 8 8 &

OOS WMS) Goounpcoaccoceosoo sence, A AO FO Bs A ws 24 21 23 22

Hind limb....................... 47 46 46 42 42 43 41 385 37 34

Nolet teehone patduscomacbcboocadou ten | CADN Bl Pe hy pes “Ost 21 19 22 18

Tail (*reproduced) ............ 170 95* 77*100* 80* 140 105 97* 98*125

1, 9. Greece; 2, 8. Olympia; 3, 6, 7. Morea; 4. Kalamata; 5, 10. L. Stymphalos.

Palatal Teeth.

These are strongly developed, forming two to four longitudinal

series on each pterygoid bone.

Scaling.

Rostral shield largely entering the nostril. Nasals forming a

short suture behind the rostral* ; frontonasal broader than long ;

Text-fig. 8.

Upper and side views of head of male and female (from photographs).

frontal not longer than frontoparietals, anterior borders very

concave, forming an acute angle, the shield often appearing

* In one specimen the frontonasal forms a short suture with the rostral.

THE PELOPONNESIAN LIZARD. 39

trilobate in front, in adult male specimens* ; parietals about once

and a half as long as broad, in contact with the upper postocular ;

occipital very variable in size, sometimes longer, sometimes

shorter, and usually broader than the interparietal, from which

it may be separated + by a short suture formed by the parietals.

Supraoculars in contact with the supraciliaries, rarely one or

two granular scales intervening between them. Postnasal single {.

Four or five upper labials anterior to the subocular $, Rather

large, irregular shields cover the temple, often exactly as in

L. viridis ; tympanic shield distinct; 3 to 5 upper temporals in

contact with the parietal. 28 to 33 scales and granules in a

straight line between the symphysis of the chin-shields and the

median collar- -plate ; gular fold distinct. Collar with even edge,

formed of 10 to 12 plates.

Scales on back juxtaposed, granular, round or suboval, feebly

or faintly keeled, on flanks towards the ventrals larger, flat and

Table showing number of scales, plates, and femoral pores.

ih) eee day One ¥1Ge 1. 8.

Bhs (CiREXED ccenaccessoescseus 0) “EY 7 GIy = ml) Si IR) Ey tas)

9 OME sconces TK ES i Bo) 4. 23 28

mee Monument naeseesces LOL Our? 288 a2 e28 4 20-21 3)

Pneealeamationweere nacre: 100) GON ZOE O30 5 21 26

x) We Sivoo bees. Wi Gay By 1K) BY) 4 22-24 8

Fy TULA) Ray et Lahr a HO G2 ey Sil SS BEDS BHI

OP Mouetemree ee eee 59 31 11 30 45 22 29

SeEOlymplaeeee ces: eid lee DOpmecaN | Il eons Fab 23 27

» Greece.. seeoheses 0) SB an BA 20 30

on We Shay mphalos SED: sy G2 SL ah, BB} 4, 24-25 27

1. Length (in millimetres) from snout to vent. 2. Number of scales across middle

of body. 3. Transverse series of ventral plates. 4. Number of plates in collar.

5. Number of scales and granules between symphysis of chin-shields and median

collar-plate. 6. Number of upper labials anterior to suhocular. 7. Number of

femoral pores. 8. Number of lamellar scales under fourth toe.

subimbricate, smooth ; 53 to 63 scales across the body, 2 or 3

transverse series corresponding to one ventral plate.

Ventral plates in 6 longitudinal and 28 to 33 transverse series.

Anal plate moderately large, with 2 or 3 semicircles of scales

around it.

‘Seales on upper surface of ies a little smaller than dorsals,

distinctly keeled; 26 to 31 lamellar scales under the fourth toe;

20 to 25 femoral pores on each side.

Caudal scales truncate behind, more or less strongly keeled, the

whorls subequal in length.

Coloration.

This varies considerably according to individuals. Females and

* In the allied Z. taurica the anterior borders of the frontal vary from feebly

concave to feebly convex, as in L. muralis and its varieties. Peracca has attached

undue weight to this character in his description of L. sardoa.

+ In 4 specimens out of 12.

{ Two regularly superposed postnasals in one specimen (2, Morea).

g 4 on both sides in five specimens 4 on one side and 5 on the other in six, 5 on

both sides in one,

AQ MR. E, G. BOULENGER ON

young are beautifully striped with dark brown or black, some

specimens may even be described as black above with 5 or 6 light

longitudinal streaks. Traces of this striation may persist in some

males, but usually disappears entirely, the back beimg uniform

brownish or dul! green, with small blackish spots or vermicular

lines on the sides. The figures annexed to this description will

give an idea of the gorgeous colours assumed by the adult male

during the breeding-season, making it one of the most beautiful

of the European Lizards. The top of the head is of a reddish

brown, sharply contrasting with the green colour of the nape,

which gradually changes to olive or brown on the posterior part

of the body; the sides of the head and body and the lower parts

are of a bright vermilion-orange, reiieved by’a patch of azure-blue

in the axillary region and a broad band of the same colour occupying

the outer row of ventral shields and extending a little way up on

the scaly part of the side. The female is of a reddish brown,

with two broad blackish bands along each side, the outer proceeding

from the eye, bordered above and below by a narrow whitish

streak and separated by a third hght streak which in some

specimens is pale yellow, in others pale green; a small round

blue spot is present above the axil; the hind limbs bear round

light spots edged with blackish; the lower parts are white or

pale yellow, often tinged with rosy or lilac on the sides.

Relationships.

Lacerta peloponnesiaca is most nearly related to Z. tawrica, and

with it fills the gap between the massive Lizards hike Z. agilis

and L. viridis and the forms that cluster round LZ. mzralis. In its

thick, convex skull and its well-developed pterygoid teeth, as well

as in its temporal scutellation, it agrees with the former group,

which I ‘regard as the most generalized, whilst in its scaling and

especially in its non-serrated collar it agrees very ciosely with the

typical L. muratis, from which some authors have held it to be

probably derived.

EXPLANATION OF PLATE f.

Lacerta peloponnesiace, male and female, natural size.

4. Remarks on Two Species of Fishes of the Genus Gobius,

from Observations made at Rosecoff. By Enwarp G.

BouLENGER *,

{Received Octoher 30, 1910: Read November 15, 1910.)

Prof. F. Guitel, in his well-known paper on the breeding-

habits of Gobias minutus ?, vemarks that the descriptions of this

* Communicated by G. A. Bourmneen. F.RS., V.P.Z.S.

7+ Arch. Zool. Exp. et Gén. x. 1892, p. 499.

FISHES OF THE GENUS GOBIUS. 4]

species by various authors vary toa considerable extent and that

the fish described under this name by Collett * does not* tally

with the said Goby of Roscoff on which he based his observations,

the number of scales along the lateral line in his specimens being

about 45, those of the Norwegian fish about 60. Further that

Collett’s formula for the fin-rays---D. 6/11-12. A. 11—differs

slightly from that of the Roscoff fish, which he gives as: D. 6/8-

10. A. 8-10.

During a stay at Roscoff last summer I collected a number of

specimens of the Goby described by Guitel under the name of

G. minutus and also found there another Gol by which differed at

a first glance from the former in its larger size, coloration, and

general appearance, and which I found on examination under a

lens to have more numerous seales. This is the form described

by Messrs. Holt and Byrnet, in their paper on the British and

Trish Gobies, as the typical G. minastes, while the fish so commonly

found at low tide in the pools of the shallow sandy bays of

Roscoff is regarded by them as an estuarine race of the same

species, to which they refer the G@. microps of Kroyer = and later

Scandinavian authors.

The colour of the latter fish is dorsally of a dirty grey, minutely

speckled with black, laterally with large blackish blotches, which

in the males usually expand into vertical bars on the side. That

of the former is creamy speckled with rusty brown and with small

blotches of the same colour laterally; the blotches may also

form bars, which, however, are always finer and less conspicuous.

J found the smaller form to be stouter than the larger, the depth

of the body being usually from 5 to 6, as against 6 to 7 times in

the total length, caudal fin excluded, and the scaleless area of

the nape and back to he of greater extent.

These two fishes I found under quite different conditions—the

G. microps of Kréyer close inshore, the one alluded to by

Messrs. Holt and Byrne as the typical G. minutus at a locality

north of the little island of Batz, opposite Rosecoff, uncovered at

the spring tides only.

From the table of particulars of the two forms, given further

on, it will be seen that they differ both in number of scales and

fin-rays ; and there can he no doubt that the two fishes are distinct

and well deserve to be regarded as valid species, not as races

only.

Messrs. Holt and Byrne, however, are of opinion that a sufficient

series of specimens from various localities would show a complete

gradation from the one “race” to the other, and state that

specimens from the Cuckmere river approach the typical form in the

large number of scales and small sealeless area of the nape and

* Vidensk. Selsk. Forh. Christiania. 1874, p. 168.

+ Report on the Sea and Inland Fisheries of Ireland for the Year 1901, Part il.

Appendix LIT. (1904).

* Damn. Fiske. 1. p. 416 (1838-1810),

49, MR. E. G. BOULENGER ON

back, though in form and colour they resemble the estuarine race.

I have examined several specimens in the British Museum from

the Cuckmere, presented by Mr. Byrne, but find that both in

appearance as well as in the number of scales they agree with

G. microps.

It seemed desirable to make sure, by reference to the original

description, which of the two species so often confounded should —

bear the name of Gobius minutus. This name was proposed by

Pallas* for the fish (“ Maris Belgici”) described in a rather

puzzling manner by Gronoviust, who gives the number of fin-

rays characteristic of G. microps, while, on the other hand, the

total length “ tres wncias” (about 80 mm.) can apply only to the

larger species. It is highly probable, however, that Gronovius

had before him examples of both species from the Belgian coast{,

and that he noted the number of fin-rays from the smaller fish

and added to his description the size attained by the larger.

Gmelin’s § diagnosis “ albicans ferrugineo maculatus, ... D. 6, 11.

A. 11” can only apply to G. minutus of most authors.

It is therefore satisfactory to find that no objection can be

raised to the retention for the two species of the names used by

the Scandinavian and other authors who have distinguished them.

The following is a tabulation of the specimens of the two

species from Roscoff || :—

G. minutus.

Length. | Scales. | Fin- Rays.

| paged)

| D.z A.

55 66 | VI. 12 11

53 | 67 ViLi2 11

53 | 65 VI. 12 12

53 | 64 VI. 12 12

50 63 VI. 12 12

| 49 71 Vi. — —

49 64 Wo Lz 11

| 48 65 VI. 12 11

| 46 63 | VI. 12 12

| 45 | 62 | VI. 12 1

Ad. | 61 Wal 11

| 39 | 63 VI. 12 12

* Spicil. Zool. vill. p. 4 (1770).

+ Zoophylacium, p. 81. no. 276 (1763).

+ Specimens of both G. minutus and G. microps from the Belgian coast were

sent to the British Museum by the late Prof. 2. van Beneden, and measurements

of these will be found in the table at the end of this paper. The two species have

hitberto been confounded by Belgian authors under the name of G. minutus.

§ Syst. Nat. i. p. 1199 (1788).

|| In these tables the length (in millimetres) is taken from the end of the snout to

the base of the caudal fin. The scales are counted in a longitudinal series from the

upper extremity of the gill-opening to the root of the caudal fin.

ee ee

FISHES OF THE GENUS GOBIUS.

G. microps.

Length. Scales. | Tin-Rays. |

| [ees ete ee is Baier OE al

D. A,

45 48 VI. 10 v0

45 50 Will's 10 '

45 } AA, Miles9 10

4A, 52 VI. 10 10

43 4d, | VI. 10 9

42 46 VI. 9 10

41 | 4.7 VI. 9 10

40 H 49 VI. 10 10

40 | 45 Wallee 9

37 40 | VI. 9 10

36 | 48 | VI. 10 10 |

34 49 VI. 10 10

In order to satisfy myself as to the degree of constancy of

these characters, I have examined a number of specimens in the

British Museum collection from various localities, including the

Cuckmere river and the Belgian coast.

The following table may be useful for purposes of comparison :—

G. minutus.

Locality. Length. | Seales. Fin- Rays. |

ee ca a ee |! met bs

D. A.

| Belgian coast (Van Beneden) ......| 53 70 VI. 12 12

2 conte | 49 Go Mains |e) ao

| Weston-super-Mare (Day) ............ | 60 70 | VI.11 11

, Plymouth (Marine Biol. Assoc.) _..| 70 71 | VI. 12 12

ae im i sr FO wis? VVelo donne:

Brishtone(Children)l sess). -| SE) WHE Tay int |

} LATDE a) ose) oS Nel ae gata Ment es peoee need AT 62 WOE aa Til

; Burford Bank, Irish Sea (Byrne) _. 57 | 61 Wiig 12

. 4 Jp ha ach eae 2) 61 Wee = ie

PL COKumn st! (SeOAne) =e eee eee 70 73 Wil, THE 11

eae NE ome a tn erty Mn) ve a0 67 VAD O11

i ve a ne 2 Femi hai ad

G@. microps.

| }

| Belgian coast (Van Beneden) ...... 43 46 VI. 10 10

ie Ait Sy as | 35 48 Miedo 10

Weston-super-Mare (Day) ............ 42 47 WAG dat 10

Cuckmere R., Sussex (Byrne)......... 43 46 VI. 10 10

= 3 5 eas Al 47 VI. 10 10

5 i (are ee 41 45 VI. 10 iit

5 op 33 eee 36 52 VI. 10 10

| 3 2p 33 eee 35 42 VI. 10 10

lpsepNorwiary: (Colle). s125) seer 38 46 VI. 9 9

| is Sa cones nee ean ee temete 33 42 | Wig 8) i)

| Denmark (Day) «..-...--1. eee 33 AAS ND lO

eae ene ero igs ti he L.VISION ANG

}

44 ON FISHES OF THE GENUS GOBIUS.

As will be seen from these tables, G. mecrops has 9 or 10,

exceptionally 11, rays in the 2nd dorsal and anal, G. minutus

having 11 or 12; the number of scales along the lateral line,

however, shows no overlap, not exceeding 52 in G. microps and

not falling below 61 in G@. minutus.

L therefore hope I have succeeded in settling the question of

the correct name of the fish so carefully described by Prof. Guitel,

about which he rightly entertained some doubts at the time of

his observations on its remarkable br eeding-habits.

The two forms here discussed have been quite correctly sepa-

rated and identified by Messrs. Holt and Byrne; the only point on

which I cannot agree with them is with regard to the existence

of connecting-links, which the examination of a large material

has failed to disclose.

In concluding this note, I wish to express my indebtedness to

Prof. Yves Delage for kindly allowing me to work at the Roscoff

Laboratory, of which he has the direction,

APPENDIX.

Col. Shepherd, who has devoted much time to the study of

otoliths, has examined for me those of the two fishes from Roscoff,

and finds their claim to specific distinction confirmed by the

ditferences in this character.

He has kindly drawn up the following notes, as an appendix to

my communication :—

“Under the microscope the otolith of Gobiws microps shows as

a quadrilateral lump with fairly equal sides. Two are at a right

angle, but the angle is rounded off; a third is bulged out anne an

amire nd curve, Te fourth forms an indent. The thr ee sides first

mentioned are plain-edged.

“The otolith of Gohiaus minatus shows an irregular quadrilateral

shape: one side is straight and plain ; the other three sides are

not so symmetrically shaped as In G. microps, and are markedly

scalloped, there being six lobes on the edges of the three sides,

these lobes not regularly spaced, but of varying size.

“This would show that the two fishes are different species.

“The otoliths referred to are in each instance the sagitta.”

ON COLOUR-PHASES OF THE BUDGERIGAR. AD

EXHIBITIONS AND NOTICES.

November 29th, 1910.

Dr. Henry Woopwarp, F.R.S., Vice-President,

in the Chair.

Dr. WiuttaAm Nicouu, of the Lister Institute of Preventive

Medicine, gave a demonst ‘ation of his method for the collection

of ilire matodes, elit

Dr. R. 'T. Lerper, F.Z.8., exhibited two photographs and

some specimens showing the Nematode infection known as

Onchocerciasis in beef imported from Queensland.

Dr. J. F. Gemuriy, M.A., D.Se., Lecturer on Embryology in the

University of Glasgow, gave an account, illustrated by lantern-

slides and specimens, of his memoir on “The Development of

Solaster endecw Forbes,” communicated to the Society by Prof.

J. Arthur Thomson, F.Z.S.

This memoir will be published entire in the Society’s ‘Trans-

actions’ in due course.

Mr. D. Seru-Smirn, F.Z.S., the Society's Curator of Birds,

exhibited living examples of the Australian Budgerigar or Un-

dulated Grass-Parrakeet (elopsittacus wndulatus), showing three

colour-phases. The normal bird was mostly green, with a yellow

face, dark barring across the occiput and back, and blue on the

tail-feathers.

The yellow variety was now common as a cage-bird, and had

been known to occur in a wild state. Init the dark pigment had

disappeared and practically all trace of blue had been eliminated,

though some spots on the cheek, which in the normal bird were

deep indigo-blue, retained a faint bluish tinge.

The third variety was an extremely rare one, in which all the

yellow pigment had gone, leaving the bird almost entirely blue.

Those parts which in the normal bird were green, were in this

variety pale blue, while the face, which was yellow in the normal

bird, was pure white.

Blue Budgerigars appeared to have been known in Belgium and

France some twenty-five or thirty years ago, as they were men-

tioned by Greene in his ‘ Parrots in Captivity’ (i. 117) and others of

his books, and by Wiener in Cassell’s ‘ Canaries and Cage Birds.’

The variety seemed to have been entirely lost sight of, however,

in this country at any rate, until M. cbmc a well-known

Belgian aviculturist, exhibited a pair at a bird-show held at the

Royal Horticultural Society’s Hall at Westminster on November

25th—28th, 1910. This eentleman had several of these birds,

which) were said to: breed true to type, but to produce a pre-

ponderance of females.

46 ON PNEUMO-ENTERITIS IN THE RED GROUSE.

Mr. Seth-Smith pointed out that, so far, no systematic breeding

experiments had been carried out with Bud gerigars, but with three

distinct colour-phases of a free-breeding species oy work with, the

material for some very interesting equeniinen ial breeding was at

hand.

He acknowledged his indebtedness to M. Pauwels for the loan

of the blue specimen exhibited at the meeting.

PAPERS.

5. Ona Possible Cause of Pneumo-enteritis in the Red Grouse

(Lagopus scoticus). By H. B. Fanruam, D.Sc., B.A.,

V.Z.S., and H. Hammonp Surra, M.R.C.S., L.R.C.P..,

ese

| Received October 24, 1910: Read November 29, 1916. ]

The importance of Coccidiosis as a serious disease of the

digestive tract of birds has lately been clearly established by

Fantham in England in the case of young Grouse and Pheasants,

and by Morse aud Hadley in America in Fowls and Turkeys.

While pursuing our researches at the Frimley Experimental

Farm belonging to the Grouse Disease Inquiry Committee during

the summer of 1910, we found that out of 40 Grouse chicks

hatched, 17 died between the ages of 4 and 6 weeks. These

birds were examined by beth of us and were found to be suffering

from Cocczidiosis, the parasites (Zimeria (Coccidiwm) avium) oceur-

ring especially in the duedenum and ceca. Many of these young

birds, however, also presented symptoms of pneumonia, consequently

the lungs, trachea, and bronchi of the birds were most carefully

exainined. The results of our examination were most interesting,

for we both found coccidian odecysts in the trachea, bronchi, and

bronchioles. Inside these odcysts the processes of formation of

the four sporoblasts were sometimes found to be going on. The

odcysts were probably acquired by the mouth, and a few of them,

instead of passing directly dewn the digestive tract, as is usual,

may have found their way, vid the glottis, into the trachea and

bronchioles. It is possible that these coccidian cysts in the

bronchioles would be quite capable of setting up sufficient irri-

tation to account for the pneumonic symptoms seen in the lungs

of these young birds. It would seem, therefore, that the old

name of pneumo-enteritis, as applied by Mr. Tegetmeier and

others to one of the diseases that caused mortality in Grouse —a

view which has met with much criticism—may after all be proved

to have some foundation in fact.

References to Literature.

Cots, L. J., and Hapuey, P. B. (1910).—* Blackhead in Turkeys,

a Study in Avian Coccidiosis.” Bulletin 141, Agric. Expt.

Station, Rhode Island State College.

Fanraam, H. B. (1910).—‘‘ On the Morphology and Life-History

of Fimeria (Coccidium) avium, a Sporozoon causing a Fatal

ON THE ALIMENTARY TRACT OF CERTAIN BIRDS. 47

Disease among young Grouse.” P. Z. 8. 1910, pp. 672-691,

4 plates.

Fanruam, H. B. (1910).—‘ Experimental Studies on Avian Cocci-

diosis, especially in relation to Grouse, Fowls, and Pigeons.”

P. Z. S$. 1910, pp. 708-722, 1 plate.

Kunin, E. (1892).—‘‘ The Etiology and Pathology of Grouse

Disease and Fowl Enteritis.”

Morsz, G. B. (1908).—‘* White Diarrhea of Chicks.” Circular

128, U.S. Dept. Agriculture, Bureau of Animal Industry.

Trcermerer, W. B.—‘ Field,’ vol. 104, p. 561 (24/9/04); vol. 105,

p. 1027 (17/6/05); vol. 106, p. 410 (26/8/05); vol. 107,

p-. £65 (24/3/06).

6. On tne Alimentary Tract of certain Birds and on the

Mesenteric Relations of the Intestinal Loops. By Franx

EK. Bepparp, M.A., F.R.S., F.Z.8., Prosector to the

Society.

[Received March 1, 1910: Read November 29, 1910. ]

(Text-figures 9-28.)

For some time past I have been accumulating notes relative to

the viscera of birds which have died in the Society’s Gardens,

and have paid special attention to the alimentary tract. In the

following pages I call attention to the intestinal tract of a

number of birds which either have not been studied or as to

which my own investigations lead me to disagree with earlier

statements.

T have dealt more particularly with such species as have not

been carefully studied from the point of view of the convolutions

of the intestine, and am able to call attention to a considerable

series of birds. ‘The subject is by no means a new one, dating as

it does from the accurate though few observations of John

Hunter. I arrive, however, at rather different classificatory

conclusions from others, and venture therefore to direct the

attention of the Society not only to the new facts but also to

certain classificatory inferences to which these facts point.

The observations which I lay before the Society may be

considered under the following headings, viz. :—

§ Historical Survey, p. 48.

§ Description of the Intestinal Tract in various Groups of

Birds, p. 50.

§ Some Generali Considerations, p. 86.

§ The Primitive Form of the Intestine in Birds, p. 86.

§ The Course of the Evolution of the Gut, p. 87.

§ The Mutual Affinities of Avian Families judged by the

Intestinal Convolutions, p. 89.

§ The Relationship between the Gut and the Nature of the

Food, p. 90.

§ Summary of Facts relating to the Intestinal Coils of

Birds, p. 92.

48 MR. F. E. BEDDARD ON THE

§ Historical Survey.

There is no doubt that the fullest general account of the

intestinal tract of birds written by the older anatomists is that of

John Hunter*. He examined and annotated upon more than

fifty species not wholly though mainly British. His observations

are entirely correct, though notalways quite fullenough. In many

‘ases, however, he has seized the main features of the intestinal

coils so accurately and sufficiently that but lttle in the way of

addition is needed. Thus in the Gallinaceous birds he has

‘appreciated the loose arrangement and absence of fixed loops in

the postduodenal section of the small intestine and the attachment

of its terminal region to the duodenum. In the Rails he has

correctly described the three distinct loops of the jejunal region

and the attachment of the first and third of these together. In

the Accipitres the short loop just above the ceca is described in

many forms. The peculiarities of the Parrot intestine are fully

described. In short, Hunter accomplished a great deal.

Cuvier and Duvernoy r distinguished perfectly correctly, as Dr.

Mitchell has pointed out, the three separate regions in the small

intestine of a bird, which I propose to call duodenal, jejunal,

and ileic loops, and they also indicated the fact that the middle of

the three loops is frequently folded upon itself, contorted into a

spiral, or subdivided into several regions. Furthermore. it is

semen ise (and I find myself im arcu with this opinion) that,

‘le canal intestinal des oiseaux est loi cde présenter des differences -

atissi nombreuses, d'une espéce, d’un genre ou d’une famille

& Vautre, que celui des mammiféres.” Thereafter follows a

considerable amount of detail concerning these different loops

in the different groups of birds. For example, the thiee simple

loops of the Passerines are referred to in a good many species and

the spiral arrangement of the middle or jejunal loop is described

in the Crows. ‘The Picarian birds, Touracou and Cuckoo, are de-

scribed in such words as to show that they agree completely with

the Passerines. Cuvier did not, however, as Dr. Mitchell has also

and quite justly pointed out, delimit the suds region correctly.

He describes the limits of the third (and last) region of the small

intestine as indicated sometimes on the side of the middle loop by

an unpaired cecum (7. ¢., Meckel’s diverticulum). This is never

the case, I believe =

In his ‘ Lectures on Comparative Anatomy,’ Sir Everard Home §

has figured the coils of the small intestine ina number of birds—

for instance, the Raven, where the spiral of the jejanum and the

close association of the ileic and duodenal loops are indicated ; the

Tats Lecons ad’ AmEiounl comparée de Georges oe ier, rec. et publ. par G. L.

Duvernoy, t. iv. 2me partie, Paris, 1835, p. 269 e¢ seq.

+ But see for a possible exception the account of the Tinamous ay p- 52.

5 ‘Lectures on Comparative Anatomy, London, 1814, vol. 1. 402, vol. ii.

pls. civ.-exii. I am indebted to Dr. Mitchell for the exact 1 fete to ‘ahr more

than once misquoted work.

ALIMENTARY TRACT OF CERTAIN BIRDS, 49)

Sea-mew, where the spiral is also shown, and it is remarked by the

author that the turns of the intestine bear a close resemblance to

those of the Crow, Swan, Goose, “ Ardea argala,” &¢.—not a very

long series of birds.

Owen, in describing the Flamingo *, pointed out that the small

intestines “‘ were disposed in twenty-one elliptical spiral con-

volutions, eleven descending towards the rectum and ten returning

towards the gizzard in the interspaces of the preceding.’ The

same anatomist correctly described the three loops in the small

intestine of the Hornbill. In the ‘Comparative Anatomy and

Physiology of Vertebrates’ many more facts are given, most of

which appear to be quite correct, but all are not quite intelligible

to myself. Not many comparisons are made. The Cuckoo

is correctly described, but it is not pointed out that it agrees

with the Hornbill, which bird, indeed, is not referred to in the

volume. ‘The general prevalence of concentric folds among birds

with long intestines is noted. The peculiarities of the Galli-

naceous birds which have no fixed loops except the duodenal are

appreciated in the description of the Common Fowl. The attach-

ment of what I term the ileic loop to the gizzard and to the

duodenal loop is mentioned.

Dr. Gadow’s contributions § to the subject of the present com-

munication have an importance of their own which is very great.

But they do not come exactly within the limits of the discussion

to which I desire here to contribute, since the aim of that

anatomist was to pourtray the arrangement of the gut within the

body-cavity and not to delimit only the permanent loops of the

intestine as formed upon the supporting mesentery.

The most recent contributions to the subject known to me are by

Dr. Chalmers Mitchell ||. In these memoirs, the author, in addition

to discussing some parts of the subject with which I am not

concerned here, deals with a much larger series of species than

any previous author and has arranged his observations syvstema-

tically, so as to cover most of the existing groups of birds. His

special object, however, was to trace the various modifications of

the intestinal tract to what he believed to be a primitive type, to

arrange them in the form of a phylogenetic tree, and to see how far

such a tree would agree with or correct conceptions of the phylo-

genetic ideas regarding birds as a whole. In the course of this

paper I shall refer to various points in which my own observa-

tions do not agree with those of Dr. Mitchell. In my opinion,

however, Dr. Mitchell’s mode of figuring the intestinal tract of

birds gives an appearance of simplicity which is misleading, with

the result that birds which are separated by marked characters

* P, Z.S. 1832, p. 142. + Ibid. 1833, p. 102.

+ Vol. ii. 1866, p. 167 ef seq.

§ “Vergl. Anatomie des Verdauungssystemes der Vigel,” Jen. Zeitschr. 1881.

“On the Taxonomic Value of the Intestinal Conyolutions in Birds,” P. Z. 8.

1889, p. 305; in Newton's ‘ Dictionary of Birds,’ sub voce “ Digestive System.”

|| “On the Intestinal Tract of Birds, &.,” Trans. Linn. Soc. viii. 1903, p. 175;

and an earlier paper in P. Z. S. 1896, p. 136.

Proc. Zoot. Soc.—1911, No. LV, 4

50 MR. F. E. BEDDARD ON THE

are represented as being almost identical. In particular, Dr.

Mitchell does not always distinguish between fixed loops, definitely

formed by a narrow mesentery, and the irregular folds into

which any mobile coil of the intestine may fall when disposed on

the dissecting-board in Dr. Mitchell’s fashion. I shall recur to

definite instances in the course of this communication.

I shall now -proceed to deal with the intestinal tract in a

number of species of birds.

§ Description of the Intestinal Tract in various

Groups of Birds.

On opening the abdominal wall of most birds the intestine is

usually seen to form a rather compacted mass, such as is figured,

for example, by Dr. Gadow in most of the plates which illustrate

his original memoir upon the intestinal tract in birds. This

mass consists of parallel or concentrically arranged loops of intes-

tine, and in the higher birds, such as a Heron, a Duck, or Stork, is

very characteristic. This appearance of the gut distinguishes it

at once from the Mammalian or Reptilian gut, where the intes-

tine lies laxly within the abdominal cavity.

This also is the case with all the Struthious birds, whose intes-

tinal tract at the first glance recalls that of a Mammal. A little

disturbance of the apparently compact intestinal mass of some

other birds, as, for instance, the Hagles and Hawks, shows that

here, too, the intestinal tract is not really much welded together,

but simply lies pushed close coil to coil, owing to the limited space

in which it has to be stowed away. In other cases, however, it

can be easily ascertained by the gentle pulling apart of the

intestinal coils that the gut is disposed in tightly fixed loops.

This is the case, for instance, with Ducks, Storks, Penguins,

and a variety of other genera and families. Inasmuch as the

lax condition of the small mtestine in such a bird as an Ostrich

recalls that of the Mammalia and Reptiles, and is really like the

intestinal tract in those Vertebrates, it is to be assumed that this

condition of the bird’s gut is the more primitive condition and

that the specialisation into definitely fixed concentrically or

parallel arranged loops, whether narrower or wider, is an index of

the higher position of the bird in the series. I shall commence the

following survey of such new facts as I have to add to the matter

in hand by dealing with the more primitive groups of birds first.

Indeed, I have not attempted in this paper to map accurately the

coils in several families of birds where they are very complicated,

such as the Stork tribe ; for I am not satisfied as to the relationship

of the coils in these birds to the more simple intestine of lower

forms. It is almost entirely with the latter that I deal in the

present communication to the Society.

Of the Ratir# I have examined all the living genera. I fully

agree with Dr. Mitchell as to the basal position in this group of

ALIMENTARY TRACT OF CERTAIN BIRDS, 51

Casuarius andl Dromceus, but I find his description defective in one

particular and the illustration which he gives of Caswarius

correspondingly inaccurate. It would be inferred from that

figure * that the gut lay in a single line without any attachment

between the ileum and duodenum; that—to use Dr. Mitchell’s

own term—there was no vestige of a supraduodenal loop. The

existence of this attachment is indicated by him in other cases by

a cut blood-vessel ; there is no such “short circuit ” represented in

his figure of the Cassowary. Nevertheless, two species of Casso-

wary which I have dissected, viz. C. australis and C. westermanni,

show such a connection, which is not, however, associated with

the formation of an ileic loop distinguishable from the jejunum.

Nor can I agree with Dr. Mitchell’s figure of Apteryx, unless,

indeed, the species examined by him (4. mantelli) differs from

that examined by myself (4. australis). For I find in the latter

bird no definite ileic loop, but only an attachment by mesentery

of the latter part of the ileum to the duodenum. The bird, in

fact, exactly resembles Casuarius, Struthio, and the Gallinaceous

birds in this particular.

In Kkhea americana the -intestine is formed upon a plan which

may be interpreted in one of two ways—one of which is certainly

not ‘‘archicentric” in the sense in which Dr. Mitchell uses the

word, and the other interpretation hardly justifies the use of the

word ‘“archicentric.” Since, in various other points of structure

(e.g. less degeneration of wing, syrinx), Rhea is much less

“Struthious” than Caswarius, it might be expected that the

intestinal tract also would be more like that of Carinate birds.

The accompanying figure (text-fig. 9, p. 52) shows the course of the

intestine in a female example of Rhea americana, and may be com-

pared with the figure drawn by Dr. Mitchell 7 from the intestinal

tract of the same species, with which I do not find myself able to

agree entirely. Dr. Mitchell, however, is perfectly right in dis-

tinguishing two loops only in the smail intestine, viz., the duodenal

and another which may or may not be the ileic loop of other birds,

or “‘supraduodenal,” as it is termed by him.

This latter loop is wider as well as longer than the duodenal

loop, and it lies parallel with it as does the ileic loop (nearly

always) in other birds, and is connected with the duodenal loop by

the usual ileo-duodenal ligament, which is long and extends nearly

to the end of the duodenal loop, while it is attached along more

than half of the length of the loop now under consideration. So

far the facts point towards the interpretation of this loop of the

small intestine in hea as being the homologue of the ileic loop

of other birds. If this interpretation be correct, then the jejunal

region or loop will be practically absent and reduced merely to the

small, tract just where the lower limb of the duodenal loop bends

round to join the lower limb of the (for the moment) alleged ileie

loop. There is, I think, nothing intrinsically absurd in this

* P. ZS. 1896, p. 140, fig. 3.

} Trans. Linn. Soe. ¢.e. p. 183, fig. 3.

52 MR. F. E. BEDDARD ON THE

suggestion; it is merely the assumption of the still further reduction

of the jejunal region of the gut which is already greatly reduced

in such birdsas Chunga burmeisteri and Houbara macqueent, where

it is already as short as or even shorter than the ileic loop. There

is another argument in favour of this interpretation of the two

well-marked intestinal loops of Ahea which is derived from a

Text-fig. 9.

Intestinal tract of Rhea americana.

Ce. Blind ends of cwca. d. Duodenal loop. 7@. Teic region. ad. Ileo-duodenal

ligament. j. Jejunal region In this and the succeeding figures the definite

loops are marked by transverse lines.

consideration of the Tinamou, Crypturus tatawpa. In the last-

mentioned bird the intestinal loops are very remarkable ; they

are represented in text-fig. 10. There is nothing in particular

to be said about the duodenal loop. This is followed by two loops,

which lie one above the other, the proximal loop lying ventrally

—

ALIMENTARY TRACT OF CERTAIN BIRDS. 53

to the more distal of the two. They are closely attached to each

other and to the duodenal by ligament and cannot be freed with-

out cutting or tearing. There is, in fact, every reason to regard

these two loops as a subdivision of the usually single ileic loop.

Moreover, the ileic loop is occasionally double in other birds; it 1s

distinctly formed of two parallel loops in Anthropoides par adisea *.

Text-fig. 10.

Intestinal tract of Crypturus tataupa.

Lettering as in text-fig. 9.

It is to be noted also that the ventrally situated of the two sub-

divisions of the presumed ileic loop is attached up to nearly its end

by ligament to the duodenal loop. There is no case known to me

among birds where the jejunal loop is thus attached.

Another argument of the same kind is to be derived from a

consideration of the intestinal tract of the Passerine Jxocinela

crassirostris. In this Bulbul, of which I have dissected only one

* Vide p. 82.

54 MR. F. E. BEDDARD ON THE

example, the tract of the small intestine consists of two loops

only, which are approximately equisized and are both rather

wide. Furthermore, the two oval loops of gut are attached to

each other along their whole length. It will be noted, therefore,

that these two loops agree in all their characteristics with those

of other Passerine and many Picarian Birds. But if this be so, it

follows that the jejunal loop in this Passerine is reduced to the

verge of disappearance. In any case, whatever be the interpre-

tation of the several regions of the small intestine in Jaxocincla

crassirostris, it seems to me to be beyond all question that there is

a very close likeness between its gut and that of Rhea, whether

the likeness be superficial and due to parallelism of development

or not.

Moreover, there is no bird known to me in which the jejunal

loop has any intimate relations through hgaments with the ileic

loop—at any rate, to anything like the degree which is exhibited

in the case of Crypturus tataupa, on the view, of course, that the

jejunal loop is represented. Finally—though naturally it is not

attempted to lay any very great stress upon this piece of evidence

—a particular relationship between the Tinamou and Rhea is by

no means an unreasonable suggestion.

There is, however, an alternative view to be taken of the intes-

tinal tract of Rhea americana. It will be observed that Meckel’s

diverticulum lies at about the middle of the lower limb of the

loop which has been provisionally regarded as the ileic loop; the

diverticulum lies nearer to the duodenum, 2. ¢. above the ends of

the two ceca. This fact would appear perhaps to militate against

the view that has just been set forth with regard to the intestinal

tract. For generally, at any rate, Meckel’s diverticulum les on

the jejunal portion of the intestine and, in fact, at about the middle

of the length of the entire small intestine.

But although this may be generally the case in birds, it 1s by no

means universally so. In Dendrocygna discolor, for example, I

find Meckel’s diverticulum to be very much nearer to the ileic loop

than to the duodenal, 7. ¢. to be not by any means in the centre

of the jejunal region. This is also clearly the case with Carpo-

coccyx radiatus as shown in Dr. Mitchell’s figure *. There is thus

no absolutely fixed position for Meckel’s diverticulum within the

jejunal region of the gut, though there are no positive facts which

lead to the inference that this diverticulum may lie within the

ileic area. If it be held that the existence of the diverticulum

fixes the jejunal region of the gut, then the intestinal tract of Ahea

is simply a slightly further development of that of Oaswarius in the

direction of the Gallinaceous birds and many Picopasseres when

there is no actual loop formed in the ileic region, but merely an

attachment by ligament to the duodenal loop.

The gut of the Ostrich has been described by Dr. Mitchell, as

well as by others. J have only some small matters to add to the

* Trans. Linn. Soc. ¢. ec. p. 243, fig. 60.

ALIMENTARY TRACT OF CERTAIN BIRDS. 5D

account by Dr. Mitchell in relation to the object of the present

paper. The duodenal lobe of the example of Struthio molybdo-

phanes has a lateral branch, as has the species examined by

Dr. Mitchell. The pancreas extends down the duodenal loop to a

point rather beyond this lateral diverticulum of the duodenal lobe.

It does not, however, by along way reach the end of the loop. It

does, however, in Apteryx. The attachment of the ileum to the

duodenal lobe is rather more marked than in Gallinaceous birds

and much more marked than in Apteryx. The ligamentum ileo-

duodenale reaches along the duodenal loop to a point beyond the

posterior termination of the pancreas in that loop. I found no

fixed loops either in the moderately long small intestine or in the

longer colon,

Text-fig. 11.

Intestinal tract of Talegalla lathami.

P. Pancreas. Other lettering as in text-fig. 9.

The GaALuinaczous birds appear to be very uniform in the

structure of the gut. I may take Crax carunculata as a type

with which the very slight divergences shown by other Galli

may be compared: the duodenal loop is long and very thick and

the pancreas extends about halfway down it. The duodenum

soon narrows to form the jejunal region, which is of considerable

length and arranged in loose folds which can be straightened

out and among which are no fixed loops. There is no sharp line

of demarcation between the jejunal and the ileic region, which

56 MR. F. E. BEDDARD ON THE

later becomes a straight tract of intestine running partly parallel

with the duodenum and attached to it just for a short space at the

root of the duodenum where it (the ileum) bends upon itself

to join the colon. A second. species, a hybrid C. globicera

and C. hecki, was absolutely identical in all the characteristics

just given. The description of one species fits the characters of

the other.

Text-fig. 12.

Intestinal tract of Ortalis ruficauda.

G. Gall-bladder. Other lettering as in text-figs. 9 & 11.

In Talegalla lathami (text-fig. 11, p. 55) the only difference that

T could detect was the furtber extension of the pancreas along the

duodenal loop, the end of which, however, it does not reach.

Among the Phasianide I have examined a few species, and

again find no differences of moment from other Gallinaceous

birds. In Thaumalea (picta and amherstic) the pancreas reaches

to quite the end of the duodenal loop, and, as in other

genera, the ileic end of the small intestine (there is, as in other

forms, no definite ileic loop) is attached to the duodenal loop by

——

ALIMENTARY TRACT OF CERTAIN BIRDS. oF

a not very long ileo-duodenal ligament. In Hupsychortya sonnini *

the intestine is shorter, but its arrangement is precisely that of

Text- fig. 13.

Intestinal tract.of Huplocamus nycthemerus, showing condition reversed

from the normal. Lettering as in text-figs. 9 & 11.

* The ceca of Hupsychortya sonnini ave remarkable in more than one way. When

the body is opened these tubes are seen to lie ina tightly closed spiral or rather

helicoid, producing at first the idea that it is the gut itself which is thus coiled.

The spiral coiling of the cca is not, however, permanent ; they can be uncoiled and

straightened with the exception of the very tip which remains coiled. Each cecum

moreover, is seen to be covered with a network of bands in which a great deal of fat

is laid down, and which forms a loosely meshed network with the long axis of the

interstices corresponding to the long axis of the cecum. Blood-vessels traverse the

strands and apparently form a corresponding network. I am disposed to compare

this with the mass of short tubular blind outgrowths from the cca in the Tinamou,

Calodromas (‘ Ibis,’ 1890, p. 61). A slight tightening of the bands referred to in

Eupsychortyx would cause a bulging of the interstitial tracts and the consequent

formation of such diyerticula,

58 MR. F. E. BEDDARD ON THE

other Gallinaceous birds, and the pancreas extends quite to the end

of the duodenal loop. Penelope superciliaris has also rather a short

intestine and the pancreas extends to the end of the duodenal

loop, thus showing that there is no distinction in this matter

between the two groups of Gallinaceous birds. Ortalis (see text-

fig. 12, p. 56) has also a very simple and short gut. I pass by a

number of other genera that I have examined and which are

Text-fig. 14.

Intestinal tract of Podargus cuvieri.

Lettering as before.

quite like those already dealt with, to consider a remarkable

variation shown by Huplocamus nycthemerus. In one specimen

the typical Gallinaceous arrangement was to be seen; the calibre

of the duodenum was much greater than that of the succeeding

part of the small intestine and the pancreas extended to the

very end of the duodenal loop. The terminal straight portion

of the ileum was attached in the usual way by ligament to the

ia a —

ALIMENTARY TRACT OF CERTAIN BIRDS. 59

duodenal loop. In another specimen (text-fig. 13, p. 57) the dis-

position of the jejunal and ileic regions was exactly reversed *.

The duodenal loop passed immediately into a straight descending

limb bent sharply upon itself at its lower extremity, and

then passed into a laxly coiled and rather long section of gut

unattached anywhere to the duodenal loop and ended eventually

in the colon. The laxly coiled region of the gut lay to the left

side instead of to the right, and there was, in fact, in this

individual an exact reverse of normal conditions.

Text-fig. 15.

Intestinal tract of Gymnorhina leuconota.

Lettering as before.

It is thus evident that the intestinal tract of the Gallinaceous

birds is very uniform throughout the group and that it is con-

stituted upon a primitive plan which is very little, if at all, in

advance of that which characterises Apterya among the Stru-

thious birds. The only difference is, indeed, that the ileo-duodenal

ligament is longer among the Gallinaceous birds—that more of

the ileum is attached to the duodenum. But this condition is

more than paralleled by Struthio, where, as already mentioned,

* J compare these later (p. 79) with the normal conditions occurring in Fratercula.

60 Mh. F. E. BEDDARD ON THE

a considerable tract of ileum is attached by the ileo-duodenal

ligament.

“The PIcartAan Birps, Cuckoos, Touracous, and PAssERINES

really form one group so far as their intestinal convolutions go. It

is impossible, as I think, to distinguish between the Picarian

Podargus cuviert* and the Passerine Gymnorhina lewconota,

which may be compared and cannot be contrasted in the accom-

panying figures (text-figs. 14, 15, pp. 58 & 59). The salient

features in these two cases appear to me to be the great width

of the ileic loop (its extreme narrowness in e.g. the Parrots places

them at the opposite end of the series) and the close mesenteric

connection by the ileo-duodenal ligament of the two loops in

question. The great width of the ileic loop in Nyctidromus,

Cypselus, and Trogon can be inferred from Dr. Mitchell's figures 7,

though he does not, except in the case of Cypselus, identify the

loop. These figures suggest undoubtedly the primitive gut of a

Ratite or Gallinaceous bird; perhaps they are comparable with

Melanerpes x.

Furthermore, the total absence of specialised loops in the

middle region of the small intestine is to be noted. Mitchell, as

well as his predecessors in this field, has commented upon the

spiral arrangement in certain Passeres , and has remarked upon

the tendency to a spiral even where there is no actual regular

spiral formation. This affects the middle or jejunal loop, and is

greater in the Raven than in any other bird which Mitchell has

described or I have examined. I found in that bird a spival of

no less than nine double turns, whereas Mitchell has figured much

fewer in Corvus capellanus. A complete spiral of this kind is,

however, not common among the Passeres. Besides the Crow

tribe I know it only in the Tanager, Huphonia violacea. The

tendency toa spiral I have observed in many Passeres, among

which I may mention a number of Birds-of-Paradise which I have

lately had the opportunity of studying: these are Diphyllodes

hunsteim, Paradisornis rudolphi, Paradisea raggiana.

It seems to be universal or nearly so for the ileo-duodenal

ligament to connect those two loops of the intestine along their

whole lengths, and also for the pancreas to extend up to the very

end of the duodenal loop. I have found both these chavacters

to exist in Jxocincla crassirostris §, Sycalis flaveola, Huphonia

violacea, Graculus religiosus, Buceros cylindrica, Turdus migra-

torius, Ptilonorhynchus violaceus, Cassidix oryxivora, and the first-

named character in a number of other genera of which I happen

to have no note as to the pancreas. Both these anatomical

features seem likely to be characteristic of the Picopasseres

generally, even if not universally found among the members of

that order of Birds.

* JT have examined two specimens of this bird.

+ Trans. Linn. Soc. fom. cit. figs. 68, 69, 70.

ae Ki infra, p. 62.

§ A peculiarity of the gut of this Passerme has been already referred to, v. p. 53.

—~

ALIMENTARY TRACT OF CERTAIN BIRDS. 61

I have not met with many divergences among the Picopasseres

from the typical structure.

One of the most abnormal types—if not the most abnormal

among the Picopasseres is the Ground Hornbill, Bucorvus abys-

sinicus. The duodenal loop is longish and the pancreas extends

nearly toits end. It is perfectly free from the ileic loop, which

is longer than it. The ileic loop, moreover, is indented at its free

extremity and thus shows signs of being bent over upon itself,

It is also considerably longer than the duodenal loop. As in

other Picopasseres, the jejunal loop is more or less divided into

two, and the distal loop of these two is attached to the outgoing

limb of the ileic loop, which on its way to the colon is looped once

in a way precisely like that shown among the Accipitres and in

some other birds. These facts are particularly interesting, because

they confirm current opinion as to the anatomical likenesses

between the Hornbills and the Hoopoe. It is plain from

Dr. Mitchell’s figure * of the intestinal tract of that bird that

Upupa epops agrees with Bucorvus in a number of the characters

to which I have referred above. He figures the two loops of the

middle part of the intestine and the small ‘“supracecal” loop,

which latter is so characteristic a feature of Bucorvus as compared

with other Picarian birds. He does not, however, advert to this

loop by that name or compare it with the “kink” which he found

in the Accipitres of both the Old and New World. Nor does he

indicate a mesenteric attachment between the jejunal and ileic loops

in Upupa such as I find in Bucorvus. It is impossible, moreover,

to be certain from Mitchell’s figure how far the ileic and duodenal

loops are connected. Their entire mutual freedom in Bucorvus is

an uncommon feature. Although Dr. Mitchell happens, as I think,

to be wrong in remarking that the character of the gut does not

unite the Hoopoes and Hornbills closely, he was perfectly right

in making that statement from the facts before him. This is

a further example of the difficulty of arriving at sound clagsi-

fieatory conclusions without an exhaustive knowledge of the

facts.

I have lately had the opportunity of examining the gut of

Upupa, and can add something to the account given by Mitchell.

It is a rather more abnormal member of the Picopasseri ine group

than I had supposed. In my specimen there was no supraceecal

kink. The duodenal loop was very wide (as Mitchell has

remarked) and rather irregular in outline at its end, suggesting,

therefore, a commencing spiral as in Gypagus and C athartes

—a fact which may be of some significance. The duodenal

loop is larger than the ileic—precisely the reverse condition

obtaining in Bucorvus. And while in Bucorvus there is no

ileo-duodenal ligament. there is a short one in Upupa not

nearly so extensive as in Picopasseres generally, and thus

bridging over the gap between Aucorvus and its allies,

* Trans, Linn. Soe. ¢.c. p. 247, fig. 65.

62 MR. F, E. BEDDARD ON THE

The jejunum has certainly two definite loops and thus,

agreeing with Bucorvus, differs from other Picopasseres. The

pancreas in a most abnormal fashion extends into the first of

these and is perhaps responsible for its formation.

It is, of course, possible that the intestinal tract of J/elanerpes

superciliaris differs from that of other Picide. But if it agree

with that of the three species reported on by Dr. Mitchell,

then I find myself in total disagreement with that writer as to

Text-fic. 16.

Intestinal tract of Melanerpes superciliaris.

Lettering as before.

the relationships of the Woodpeckers. He observes of the Picide

that ‘the conformation of the gut is in every important respect

similar to that found in Megalema.” I have not dissected

Megalema for the purposes of the present communication, but

I have examined three species of Toucans, of which family

(Rhamphastide) Dr. Mitchell remarks that “ Meckel’s tract

and the short rectum do not differ from the form found in

Megalema.” Now in Megaleme asiatica, as is plainly shown in

ALIMENTARY TRACT OF CERTAIN BIRDS. 63

Dr. Mitchell’s figure *, there is a distinct ileic loop, which is wide

as in Picopasseres generally. In the Toucans Aulacorhamphus

sulcatus, Rhamphastos ariel, kh. carinatus, there is also a perfectly

distinct ileic loop a little less distinct than in some Picopasseres,

but still distinct.

In Melanerpes superciliaris the only properly marked intestinal

loop is the duodenal, down to the very end of which extends

the pancreas. There is absolutely no ileic loop, as is shown in

the figure (text-fig. 16). The jejunum simply passes forward

and is connected by ligaments with the gizzard and with the

duodenum quite far from the free end of the loop; it then turns

abruptly backwards, passing straight to the cloaca.

The arrangement of the gut is, in fact, precisely that of the

Gallinaceous birds, though, of course, the gut is shorter than that

of most.

Text-fig. 17.

Intestinal tract of Gecinus viridis.

Lettering as before.

There is, indeed, no great disparity in length between the gut

of Melanerpes and that of an equisized Gallinaceous bird, such

as Coturnix chinensis. In view of the primitive nature of the

palate as urged by Huxley and Parker, though not held by some

* Trans. Linn. Soe. tom. cit. fig. 71, p. 253.

64 MR. F. E, BEDDARD ON THE

others, the existence of a primitive type of gut is not without

interest. There is in any case no doubt about their difference

in this respect from their supposed allies the Rhamphastide,

though it remains to be seen whether they are like Picarian

birds of any other groups. These facts and considerations gain

additional significance from the quite similar intestinal tract of

Gecinus viridis (see text-fig. 17, p. 63).

Opinions undoubtedly differ as to the generic subdivisions of the

family Alcedinide, but Alcedo ispida and Halcyon sancta have

been placed in different genera—whether Halcyon or Sawropatis.

The intestinal tract is, however, rather different in these two

species, though one may be considered’ to be an exaggeration of

the other. The simpler of the two is that of Alcedo ispida.

In this Kingfisher the duodenal loop is free from the ileic for

at least the greater part, and thus contrasts with most other

Picarian birds. The jejunal region lies in a short spiral; but

this spiral is not a permanent structure. It can be easily

disarranged and spread out into an irregularly shaped loop.

There is, in fact, no mesenteric connection between the circles of

the spiral. The ileic loop is large, wide, and somewhat irregular,

fully as long as the duodenal loop. The spiral of the jejunal

region is, it should be added, quite a short one with only two

complete turns.

In Halcyon sancta there are differences in nearly all of these

features. The duodenal loop is, however, the same; it is a simple

loop, not particularly wide, and the pancreas extends along it

quite to its free end. I omitted tomake any notes about the

pancreas of H. vagans. The jejunal region of Halcyon sancta forms

a spiral of eight limbs, and is thus, in the first place, much more

complex than that of Alcedo ispida. In the second place, this

spiral is fixed, and is a perfectly permanent structure which

cannot be unwrapped without tearing the connecting sheets of

mesentery. These are two important differences from the spiral

found in <Alcedo ispida and are, indeed, much greater differences

than are known to me to exist between two species of any other

genus. The condition of the jejunal section of the small

intestine does not, however, exhaust the differences which even-

tually distinguish these two species of Kingfishers.

The ileic loop is, as in the last species, quite free from the

duodenal; there is no ileo-duodenal ligament, except perhaps at -

the very base of the otherwise mutually free loops. The loop

is, however, double, as it is, for example, in Grus japonicus *,

and as is shown in text-figure 18. Of these two loops, the

proximal is the larger and is wide and somewhat irregular in

form, and of about the same length as the duodenal. On the

whole, it may, as I think, be admitted that the difference which

the alimentary tract of this Picarian bird shows from that of

other Picarian birds is actually greater than that which exists

* Vide p. 82.

ALIMENTARY TRACT OF CERTAIN BIRDS. 65

between the Limicoline Recwrvirostra and such a Passerine as

Euphonia—in this case, two entirely different groups of birds.

Among the CucutrI have examined only Hudynamis orientalis,

which is one of the genera which does not seem to have been

examined by previous writers from the present point of view.

Text-fig. 18.

Tntestinal tract of Grws japonicus.

Lettering as before.

So far as Iam able to say from the examination of this one type,

Miss Marshall’s figure * of Geococcyx californianus is a better

representation of the characters of this group than that given by

fig. 22.

Proc, Zoo, Soc.—1911, No. Ve 5

66 MR. F. E. BEDDARD ON THE

Dr. Mitchell *. For the jejunal region possesses no definite loops

and the gut, as a whole, is entirely like that of the majority of

the Picopasseres. In Hudynamis orientalis the pancreas extends

to the very end of the duodenal loop; the ileo-duodenal ligament

also extends in its attachment to nearly the end of the duodenal

loop. The ileic loop is, however, longer than the duodenal.

Tn the jejunal region there are no specialised loops, and this part

of the gut shows indications of a spiral arrangement.

Of the Musophagide I have examined two examples of

Turacus macrorhynchus and one of TZ. buffoni. The two

individuals of 7’. macrorhynchus agreed absolutely ; the gut of

T. buffoni differed very slightly from that of its congener. In

Turacus macrorhynchus the duodenal and ileic loops were closely

connected throughout their whole length by an ileo-duodenal

ligament. The loops were moderately wide and of equal length

or very nearly so. The jejunal loop is a little longer than either

of the others and has a slight tendency to a spiral; when

straightened out forcibly it lies in a Y shape with some slight

rotation, as Hunter has figured in the case of Scythrops nove-

hollandie +. The pancreas, it should be observed, extends down

to the very end of the duodenal loop. The only difference that

I could detect in Turacus buffoni is that the ileic loop is rather

longer than the duodenal. It is clear that the intestinal tract

of these birds is precisely like that of the Cuckoos and of the

majority of the Picopasseres.

Of the group Hemipopi or Turnices I have dissected two

examples of the species 7'urnix varia. They were quite identical

in the convolutions of the intestinal tract. The duodenal and

the ileic loops were attached up to the end or very nearly so by an

ileo-duodenal ligament. The loops were also fairly broad. The

jejunal region of the gut lying between these two loops at either

extremity of the canal was formed of a single loop, which had a

tendency to twist itself into apparently two loops; but, without

tearing or in any way interferig with the mesentery, this part

of the gut could be moulded into the characteristic Passerine

plan, as is shown in the accompanying figure (text-fig. 19). The

pancreas extends right to the end of the duodenal loop. It is

obvious from what has been said that this bird has a typically

Passerine gut. It has not the faintest likeness to any Gallinaceous

bird. Its likeness to many Passerines is shown by the fact that

the pancreas extends down to the very end of the duodenal loop,

and also by the breadth and connection up to the very end or

nearly so of the ileic and duodenal loops as well as by the slightly

spiral, and limited, jejunal loop.

AccrpirrEs.—Among the Accipitrine birds which have not been

examined by Dr. Mitchell I have dissected the Harpy Hagle, Harpy-

haliaétus coronatus. The duodenal loop is moderately wide and the

pancreas does not extend fardown it. The ileic loop is also fairly

* Toc. cit. p. 242, fig. 60.

+ ‘Essays and Observations,’ yol. 11. p. 286.

ALIMENTARY TRACT OF CERTAIN BIRDS. 67

broad, and contrasts very markedly with the duodenal loop, on

account of the small calibre of the intestinal canal in this region ;

the duodenum is particularly wide and tapers off gradually

towards the jejunum, the greater part of which is also very

narrow; the tube is, in fact, quite as narrow here as is the ileum.

The contrast between the duodenum and the commencement of

the jejunum on the one hand, and the rest of the jejunum and the

ileum on the other hand, is remarkably like that seen in the

Gallinaceous birds, e. g., Craw (see p. 55). The ileic loop isas long

as the duodenal loop or very nearly so. It is connected to it by a

very short ileo-duodenal ligament, which leaves almost the whole of

Text-fig. 19.

Intestinal tract of Turnix varia.

Lettering as before.

both loops free of each other. The jejunal region is of considerable

length, and lies loosely and irregularly folded in the body-cavity for

the most part. The commencement of this part of the intestine,

however, where it joins the duodenum is not only, as already

mentioned, of greater calibre than the rest, but is fixed in a short

and wide and therefore not very well-marked loop. The rest of

this section of the intestine lies loosely like the Mammalian small

intestine or the jejunum in Gallinaceous birds—that is to say, it

has no fixed loops, but can be passed in a straight line between

the fingers without tearing or distorting the mesentery which

: Be

68 MR. F. E. BEDDARD ON THE

supportsit. Finally, between the ileic loop and the colon is a well-

marked supraceecal fold, found in all Accipitrine birds and in some

others. Of other Falconidee I have examined the following species,

which are not referred to in the memoir by Dr. Mitchell,

viz. Geranoaétus melanoleucus, Astur palumbarius, Tinnuneculus

dlaudarius, Astur approximans, as well as one or two species that

are referred to by him.

In Astur palumbarius and A. approximans the gut shows no

marked differences from that of Harpyhaliaétus, which I have

taken as the type for this group. The pancreas extends but a short

way along the duodenal loop, and the ileic loop is attached to the

duodenal by a short ligament only at the base. The supracecal

loop is present, and the jejunum cannot be said to possess any

pronounced folds independent of each other. As it lay in the

body I noticed the formation of spirals in this region of the gut

in A. palumbarius; but these were not permanent formations

as in Lecurvirostra avocetta. 'The whole of the jejunum could be

straightened out bit by bit, the most pronounced fold, close to the

duodenum, being large and wide and hardly comparable to the

definite folds in the jejunum of more specialised birds, such as

the Psittaci. Geranoaétus melanoleucus shows again no salient

differences ; the supracecal fold, however, is nearly as long as the

ileic loop.

I agree with Dr. Mitchell in regarding the gut of Falco

as being aberrant when compared with that of other Hawks and

Eagles. The duodenal loop is, as he has said, irregular in form.

I may add that the pancreas extends a good deal further down the

duodenal than in the other Accipitres hitherto dealt with in the

present communication. Even the ileo-duodenal ligament is a

little more extensive than it is in Astwr &e.

In Tinnunculus alaudarius there is an exaggeration of the

“abnormality ” of the duodenal lobe, which is almost bent upon

itself in a spiral fashion. In this Hawk the greater part of the

jejunum is disposed in a temporary spiral coil; but the first part

of the jejunum is in the form of a single loop, which is comparable

to that figured by Mitchell in Falco *, and which i have described

above in Harpyhaliaéius.

Spizaétus bellicosus (see text-fig. 20)is another species upon which

Dr. Mitchell had not the opportunity of reporting. It agrees with

other Accipitres in its general characters, but there are some minor

points of difference. Thus, the disposal of the jejunal is exactly

what we find in Harpyhaliaéius coronatus. This region of the

gut commences with a very wide stiff loop and then passes into a

loosely folded length of tube. The ileic loop is rather longer than

the duodenal, and the supracecal kink is developed into a loop

nearly as long, the two together reminding us of the double ileic

loop of the Cranes and even the Tinamous.

* In both of two examples of Falco peregrinus I have not seen a marked Joop

corresponding to this. The jejunum lay entirely or mostly in a rough spiral, which

could be arranged in an irregular circular fold.

ALIMENTARY TRACT OF CERTAIN BIRDS. 69

I have dissected one example of the New World Vulture,

Gypagus papa, whose intestinal tract Iam able to compare with

that of Cathartes aura described by Mitchell. In view of the

fact that the American Vultures have been regarded by many

systematists as being quite remote in their affinities from the Old

World Vultures, indeed from the Accipitres generally, it is

very important to bring together all contributions that are possible

towards settling this vexed question. The duodenal loop differs

from that of Cathartes and is, indeed, quite like that of Falco

feldeggi figured by Mitchell*. The distal extremity is folded over

Text-fig. 20.

Intestinal tract of Spizaétus bellicosus.

Lettering as before.

upon itself in a fashion that does not appear to occur among

Picarian birds. The jejunal loop is arranged in a spiral fashion like

many birds, including, however, Zinnunculus, in which Gypagus

appears to differ from Cathartes. The ileic loop is simple. The

kink so characteristic of Accipitres (but also found in other birds,

including Bucorvus) above the position of the ceca in other birds

is present as in Cathartes. It is not, in fact, possible to locate

Gypagus definitely in the system.

I have also examined Cathartes aura (see text-fig. 21), and in

most matters I am able to confirm Mitchell, as will have been

* Trans, Linn. Soc. ¢. ¢. fig. 38, p. 211,

70 MR. F. E. BEDDARD ON THE

inferred from what has been already said about Gypagus. ‘The

spiral of the duodenum makes rather more than a complete circle

and its limbs are fixed together by mesenteries; the gut does not

simply lie in a spiral. The loop which is lettered “7” in Mitchell’s

figure is broader and not so narrow according to my observations,

but better marked than in Gypagus. As to the following portion

of the small intestine, I do not agree in detail with Mitchell.

Text-fig. 21.

Intestinal tract of Cathartes aura.

Lettering as before.

The two loops forming it run close together, so that it is long

and narrow, and not broad as figured by Mitchell. There is a

tendency to form a rough kind of spiral not nearly so marked as

in Gypagus. ‘The ileic loop is only attached to the duodenal by

mesentery at its very base. The supracecal loop is more marked

than in Gypagus.

ALIMENTARY TRACT OF CERTAIN BIRDS. 71

Strices.— Among the Owls, I have examined Scotopelia bouviert,

which has not yet been investigated from the present point of

view. The duodenal loop is widish and there is no trace of an

ileo-duodenal ligament—in fact, the ileic loop lay on the right side

of the body. The ileic loop is quite simple and isabout as long as

the duodenal. The jejunum shows three quite distinct loops ;

the first of these, 2. e. that immediately following upon the

duodenum, is wider than, but of about the same length as,

another short loop which immediately follows it. A third loop of

greater length hasa distinct hint of spiral twisting. In Syrnium

aluco, Asio otus (see text-fig. 22), Strix perlata, Ninox boobook,

Bubo maximus, B. virginianus, B. maculosus, B. cinerascens, and

Strix flammea, the ileic and duodenal loops are connected by a

ligament which extends about halfway along the former loop.

The difference is rather extraordinary in the matter of this

ligament between Scotopelia and other genera, and it is, of

course, possible that we have to do with an individual variation

of Scotopelia bouvieri.

Text-fig, 22.

Intestinal tract of Asio ofus.

Lettering as before.

The division of the jejunal tract of the small intestine into

separate loops is not always well marked. In Bubo maculosus, for

example, there are no fixed folds whatever between the duodenal

and ileic loops; the whole of the jejunal region is like that of

the Gallinaceous birds or the Mammalia, and can be passed through

the fingers in a straight line without rupturing or even straining

the mesentery. The same statement applies to Bubo cinerascens, of

which species I have dissected two examples. J noticed here that

the undisturbed jejunum lay in slight spiral coils; but these were

os MR. F. E. BEDDARD ON THE

in no way permanent structures, but could be readily pulled out

straight. In Bubo virginianus the greater part of the jejunum

hes in the same way in loose movable coils; but there is a hint

of a fixed loop—wide and shallow—at the commencement of the

jejunum. Bubo capensis (I have seen two examples) and

B. maximus were quite like B. maculosus and B. ciner Uscens, a

slight and quite unfixed spiral being particularly noticeable in the

case of the first-mentioned species.

I quite agree with Dr. Mitchell that Stria flammea has a

jejunum which may be regarded as archicentric. JI cannot,

however, understand why Dr. Mitchell should emphasise the

archaic charactor of the gut of this Owl by bevmuing) it ‘‘remark-

ably archicentric” and by figuring a state of affairs * which is not

at all archicentric. For, in his figure of Strix flammea, there is

correctly represented a well-marked ‘‘supraduodenal fold ”—or

ileic loop, as I prefer to call it—and a definite loop in the jejunal

region. The latter I did not find in that species of Strix; but I

am not at all disposed to dispute the accuracy of Dr. Mitchell’s

figure. For in Strix perlatay the jejunal fold is disposed in

three more or less equisized and not very close loops; but still

they appear to be definite loops, and the jejunum is not merely

a loose coil as in Bubo. This species is therefore not at all

archicentric.

The conditions seen in Strix perlata seem to me to be a slight

exaggeration of those which J noticed in Séria punctatissima.

Jn the latter Owl the jejunum is formed by a tube which lies

in the undisturbed intestine as a spiral. It can be smoothed

out without tearing any mesenteric connections into the not

circular but rather W-shaped coil so characteristic of Passerine

and many Picarian birds; the rest of the small intestine is of

less calibre and becomes suddenly so; it again lies in the body

in a spiral fashion, but can be smoothed out in the same way

into a broad but rather irregular ileic loop. The ligamentum ileo-

duodenale extends nearly to the end of this and is attached to

about halfway down the duodenal loop. The pancreas extends

for rather more than halfway down the duodenal loop. The

gut of this genus is considerably shorter than in, for instance,

Bubo, and is to be contrasted by its stiffness with ae lower coils

of the latter.

Athene noctua, being a small species, might be expected to show

those differences from other Owls which are often met with in

comparing small species with larger allies. As a matter of fact,

the jejunal region of the gut is comparatively short and shows

no trace of any fixed loop such as occurs in some other Owls.

In this species the pancreas extends to the very end of the

duodenal loop; in a species of Ciccaha, in Strix flammea, Asio

otus, Ninox boobook, Bubo maculosus, B. cinerascens, B. virgini-

anus, and some other Owls, the pancreas does not extend so far

* Trans. Linn. Soe. ¢. ¢. fig. 66, p. 248.

+ It is not certain how far these alleged species of Strix have that value.

ALIMENTARY TRACT OF CERTAIN, BIRDS, 73

down the duodenal loop, but it is longer than in the Accipitres.

This fact, indeed, and the rather greater extension of the

ligamentum ileo-duodenale, is the chief difference that distin-

guishes the Owls from the Accipitres, the general plan of the gut

in these two subdivisions of birds being otherwise similar in many

ways.

The groups that have been hitherto considered, viz. the Ratite,

Galli, Hemipodii, Picopasseres, Cuculi, Musophagi, Accipitres, and

Striges, agree with each other in that the jejunal region of the gut,

though it may vary greatly in length, is never thrown into much

marked fixed loops, such as those which characterise the groups of

birds that remain to be dealt with. There is, indeed, the commence-

ment of the formation of such loops to be seen in the Accipitres

and Striges; but they do not arrive at the perfection and com-

plexity of interconnection which is exhibited in the remaining

families of Birds. Among the latter, however, with which I shall

proceed immediately to deal, there are species and even genera

which show the simpler conditions of the jejunum that characterise

the families of Birds already dealt with—tfor instance, in Pluvianus

among the Limicole and in the Bustards and Cariamide. In the

latter the simple conditions look like reduction ; while in Pluvianus

we may have to deal with an archaic representative of its family

which has not yet cast off the comparatively primitive type of

gut,

AuectoripEs.—The Bustards are an example of a well-marked

family of birds which show a great uniformity in their intestinal

tract. The species which I have myself examined are Hupodotis

australis and Houbara macqueent, and they evidently agree with

Otis tarda as described by Mitchell*. In Houbara macqueeni the

duodenal loop is attached to the ileie by a ligament which extends

to the very end of the former; the ileic loop is considerably

longer than the duodenal. The pancreas extends as far as the

very end of the duodenal loop. ‘T'he jejunal region is formed of a

single fixed loop, which is not quite so narrow as is depicted in

Otis tarda. Eupodotis australis (see text-fig. 23) has an intestine

which is so like that of Houbara that I can find no fresh terms in

which to describe it. Of birds admitted to be possibly allied to the

Bustards, that which most closely resembles the two genera just

referred to is Chunga burmeistert. The resemblance, however,

does not quite reach, though it very nearly approaches, identity.

In this bird the duodenal loop is, as in the Bustards, shorter than

the ileic. But the ileo-duodenal ligament stops about halfway

along the duodenal loop, though extending further along the ileic.

A point of likeness to the Bustards is the extension of the

pancreas to the end of the duodenal loop. The jejunal region of

the gut is also like that of the Bustards, in that it consists of but

one loop which occupies the whole region, of which, in fact, this

section of the intestine solely consists. Here we have an obvious

* Trans. Linn. Soc. é. c. p. 226, fig. 45.

74 MR. F. E. BEDDARD ON THE

likeness to the Bustards. The loop differs, however, in being

considerably wider than it is in the Bustards. These birds form

together a part of Mr. Sclater’s Order Alectorides, and, as they

obviously agree together very closely in the characters of the gut,

I deal with these provisionally under that name ; for there is, at

any rate, no very general agreement as to their position among

related groups and their affinities with each other.

Text-fig. 25.

Intestinal tract of Hupodotis australis.

Lettering as before.

Among the Lricota, with which, as I think, the Gulls and

Terns are obviously to be placed, there are several variations to

be seen in the coils of the alimentary tract. The most primitive

form of the alimentary tract known to me among those birds is

shown in the case of Pluvianus egyptius, for reasons which I

shall indicate after describing the facts. The duodenal loop is

fairly wide and the pancreas extends back to the very. end of that

loop. The jejunum is not definitely distinguishable from the

ileum, but the whole length of the small intestine, before it bends

upon itself to form the straight region which bears the small

and Passerine ceca, is loosely disposed as in Gallinaceous birds.

The last part of this jejuno-ileic region runs, as in Gallinaceous

ALIMENTARY ‘TRACT OF CERTAIN BIRDS, 75

birds, parallel with the duodenal loop and is attached to it by an

ileo-duodenal ligament, which extends nearly to the end of the

duodenal loop. It is, as I think, possible to interpret this intes-

tinal tract in only one way, and that is as follows.

It has been compared with that ofa Gallinaceous bird, and this

is really tantamount to saying that in the coils of the intestinal

tract Pluvianws presents us with archaic characters. After the

duodenal loop there is no marked differentiation of the gut

Text-fig. 24.

Intestinal tract of Pluvianus egyptius.

Lettering as before.

mto special loops at all. There is, however, as it appears to me,

an indication of an advance upon the condition of the gut which

characterises the Gallinaceous birds and in the direction of some

other Limicolous birds. In the Gallinaceous birds the distal

extremity of the small intestine is straightened out, but it is

relatively only a small part of the jejuno-ileum which is thus

76 MR. F. E. BEDDARD ON THE

differentiated from the much longer coiled region. Now, in

Pluvianus the distal portion of the jejuno-ileum can be de rigeur

divided off from the point lettered a in the figure (text-fig. 24),

after which point the intestine exhibits no resting in loose folds,

but passes in a broad curve to the point where it is attached

by a ligament to the duodenum. We have, in fact, here the

commencing separation from the jejunal region of a very wide

ileic loop. There is, in fact, a close similarity with the gut

of Melanerpes*. A slight alteration in the gut of both of these

Text fig. 25.

Intestinal tract of Gdienemus scolopax.

Lettering as before.

birds leads to that of many Picopasseres, where the ileic loop is

more definitely marked off from the jejunal but remains very

broad. From this type, moreover, can be readily deduced the

plan of intestinal coiling which is found in some other Limi-

colous birds which I have examined. One of the simplest of these

is Sarciophorus pectoralis, in which the ileic loop is attached to

the duodenal for nearly its whole length by the usual ligament

and is also a wide loop. ‘The jejunal has no fixed loops, but lies

* Vide p. 62.

ALIMENTARY TRACT OF CERTAIN BIRDS. id

in rather stiff coils, which approach a spiral; there are about three

of these, so that the gut is not long. It is quite difficult to

differentiate this gut from that of most Picopasseres. CU/dicnemus

(@. scolopax and (. grallarius) hardly differs from Sarciophorus.

Of this genus I may, in the first place, remark that the pancreas

extends back to the very end of the duodenal loop. The attachment

of this loop to the ileic is as in Pluvianus. The ileic loop, more-

over, at any rate in Wdicnemus grallarius, is very wide, another

point of likeness to Pluvianus and, indeed, to other Limicolous

birds that will be mentioned presently; indeed, in WW. scolopax

(see text-fig. 25) a separate ileic Joop can hardly be defined—this

character, in fact, is of the group. In both species of the genus

Cdicnemus that I have examined the jejunum lies more or less

in a spiral, which is most marked in W@. grallarius, though it is

only a short spiral of one complete turn even in that species.

It is particularly to be noted that this spiral, like that of Passerine

birds, is not a permanent spiral, but that it can be pulled out to

form an irregular circle without tearing any mesenteries. The

genera which have just been dealt with are, in fact, not far

removed from the common ground-plan, and the steps of differenti-

ation are quite as is foundin the great division of the Picopasseres.

A further stage of differentiation is seen in Securvirostra

avocetta. Dr. Mitchell has correctly commented upon the spiral

formation of the middle part of the gut, the jejunal region of the

nomenclature adopted in the present paper.

This bird shows the typical Limicoline characters in (a) the

fact that the pancreas extends to the very end of the duodenal

loop, (6) in the wide ileic loop, which is about as long as the

duodenal, and (c) in the extent of the ileo-duodenalligament. The

spiral is a fairly regular one, and although certainly not longer than,

and, I think, hardly as long as, that of the Raven, differs from it in

the important fact that it is a permanent spiral. Theseveral coils

are, indeed, connected together by mesentery and cannot be

separated out without tearing this mesentery. What is a tempo-

rary character in the more archaic forms of gut has here become a

permanent feature.

The Lari of Dr. Gadow’s classification, which I myself prefer

to associate more closely with the Limicolous birds, have an

intestinal tract which entirely justifies the latter placing. I have

examined Larus ridibundus and L. argentatus among the Gulls.

In L. ridibundus the pancreas, as i Limicolous birds, extends to

the very end of the duodenal loop. The ileic loop is also wide

and is attached for the greater part of its length by the ileo-

duodenal ligament to rather more than the first half of the

duodenal loop. The whole of the jejunum, which is rather long,

lies in loose folds lke the Mammalian small intestine, with no

fixed loops at all. I could not see any trace of a spiral arrange-

ment in this specimen. In ZL. argentatus, however, the jejunum

lay ina biggish loose spiral, which was not in any way permanent.

There is, in fact, no difference between these two species of Larus.

78 MR. F. E. BEDDARD ON THE

In both, it should be added, there was a distinet supracecal

kink.

I have dissected two examples of Sterna fluviatilis which agree

absolutely in the characters of their gut. This genus—also like

Larus—is very definitely like the Limicolous birds. The pancreas,

as in those birds, extends back to the end of the duodenal loop.

The ileic loop is wide and of about the same length as the duo-

denal and, finally, the ileo-duodenal ligament is extensive and

reaches nearly to the end of the ileic loop in the one case, and for

more than halfway along the duodenum in the other. The

jejunum lies in a rather short spiral of not more than three

circles; the coils of the spiral, however, are not so fixed as in

Recurvirostra, they can be pulled apart and arranged in about

three loops. This state of affairs has been figured by Dr. Mitchell

for Sterna hirundo, and his figure would fit perfectly the con-

ditions which I found to characterise Sterna fluviatilis. There is

also a supracecal kink.

The Auks are associated by some with the Gulls*, but by

others they are regarded as forming a distinct assemblage 7 or are

associated with the Grebes and Divers into one group Pygopodes f.

An examination of the intestinal tract of /ratercula arctica (text-

fig. 26) leads me to reject the former view and to hesitate between

one or other of the two latter classificatory schemes. At the same

time, it must be added that there is room for divergence of opinion

in the interpretation of certain of the loops, as will be seen from

the following account, in which I am unable wholly to confirm

Dr. Mitchell’s account. The duodenal loop is not in any way

remarkable and quite unfolded. The pancreas extends nearly,

or in one specimen quite, to its end. Thereafter follow two loops,

which belong to the middle (jejunal) region of the gut. These

are set more or less at right angles to each other; but the

direction of the loop is, as I think, of less importance than the

fact that there are two of them (and two only), which are roughly

equal in size and very distinct.

Moreover, these two loops are interconnected by mesenteries.

It is obvious that we have here a close resemblance to the Grebes,

Tachybaptes and Podiceps §, and to those birds only among those

whose anatomy in this respect is known. After these follows

a, large ileic loop, which is different in structure. to that of many

birds. Mitchell figures it as a simple wide loop. It is, however,

long and irregularly looped, and longer than the duodenal loop.

It appeared to me that it ended in two prolongations at its blind

end, in which case there is an obvious comparison possible with

the ileic loop in Podiceps cristatus ||, which is loosely folded at its

blind extremity. The small intestine before the ceca is thrown

into another loop, which Dr. Mitchell has duly noted and has

* H.g., Mitchell, loc. cit.

+ E.g., Beddard, ‘The Structure and Classification of Birds’: London, 1898.

t EH. g., Vertebrate List Zool. Soc. Lond. 1896.

§ See below, p. 81. || See p, 81.

ALIMENTARY TRACT OF CERTAIN BIRDS. 79

identified with that short loop whieh he has termed the “ supra-

cecal kink.” In a second specimen, all these characters were

quite as plain, so that it cannot be held that the first individual

was in any way abnormal. Probability, at any rate, indicates this

conclusion. A third example, a quite young and immature bird,

presented some slight differences which led me at first to suspect

Text-fig. 26.

Intestinal tract of Pratercala arctica.

Lettering as before.

a non-identity of species. I am assured, however, that this

suspicion is wrong. The only difference concerned the two

jejunal loops. Of these the first, 2. e. that immediately following

the duodenum, is much shorter than the second or more distal

loop. The latter, instead of being a simple loop equisized

with the first loop, is much longer and has a kink, or sudden flexure

80 MR. F. E. BEDDARD ON THE

to one side, in the middle of its course. It looks, in fact, as if two

originally distinct loops were in process of reduction to one. The

consideration of this specimen, therefore, renders another com-

parison possible, namely with the Cranes *, for in these birds the

jejunum has three separate loops. In other respects, the third

example of /ratercula arctica agreed entirely with the other two.

Apart from possible resemblances to other groups of birds that

have been indicated, the gut of Hratercula shows a feature of

particular interest in the great length and irregular disposition of

its ileic region. This latter is quite unattached to the duodenal

loop, except perhaps at the very base; it is long and lies loosely

coiled like the Mammalan small intestine. It is, in fact, the

longest section of the gut. This bird, in fact, shows an intestine

constructed in a way which is exactly opposite to that which is

prevalent in the class Aves. When there are tracts of primitive

undifferentiated small intestine left it is nearly always the jejunum

that 1s involved, and not the ileic region. In Fratercula the

jejunum is specialised imto fixed loops, while the ileum has

remained unspecialised. Indeed, my experience of this structure

among birds has only furnished one example at all parallel to ite

gut of Fratercula arctica,

This was an example of the Pheasant, Huplocamus Le hire merns

The Gallinaceous birds (see p. 55) are very uniform in the

disposition of their intestinal tract, and one out of two examples

of this species which I have dissected was perfectly normal im the

structure of the gut. A second individual, however, differed.

The duodenal loop was immediately followed by a straight, stifily

fixed, descending tube of intestine, which bent back upon itself for

a short distance and then passed into a long loosely coiled region,

a kink became continuous ultimately with the straight portion

of the ileum running to the junction with the ceca. This coiled

region of the gut, although lying on the left side of the body, was

not in any way attached to the duodenum. ‘There is, therefore,

here, it will be seen, a reversal of the conditions found generally

among Gallinaceous birds. In this example of Huplocamus

nycthemerus the stiff descending intestine which immediately

follows the duodenum obviously represents in one sense the

equally stiff ascending piece of intestine which is in the other

example of H. nycthemerus, and also in other Gallinaceous birds,

parallel to and partly fixed by ligament to the duodenal loop ;

while in the latter the loose coils which form the major part of

the small intestine lie to the right hand. There is, in fact, a pre-

cise reversal of the “normal” conditions. There is, as I think, an

undoubted resemblance between this ‘‘abnormal” example of

Euplocamus nycthemerus and the normal arrangement of the

intestinal tract in Yratercula arctica.

PopicrpEDES.—I cannot quite explain by means of Dr. Mitchell’s

figures the intestinal loops of the two Grebes TZachybaptes

* Vide p. 82.

ALIMENTARY TRACT OF CERTAIN BIRDS. 8]

JSluviatilis * and Podiceps cristatus T, of the former of which species

I have examined two examples. The plan of the intestinal tract

in the Grebes is very distinct and unlike that of any other birds

that I have studied. In Zachybaptes the duodenal loop is con-

siderably longer than the ileic (the precise reverse of the conditions

obtaining inthe Parrots), and the two are attached by a duodeno-

ileic ligament which extends about halfway along the ileic loop

(see text-fig. 27). I find only two jejunal loops (Mitchell figures

four—two long and two short, exactly as in Ara ararauna), which

are long and closely adherent for the whole of their length. The

inner of the two loops, that which is immediately connected with

the ileic loop, bore in one specimen a Meckel’s diverticulum, as

is indeed figured by Mitchell. The exca extend about halfway

along the ileic loop.

Text-fig, 27.

Intestinal tract of Tachybaptes fluviatilis.

Lettering as before.

Podiceps cristatus conforms to the general plan seen in the last

species, but differs in detail. The ileic loop is peculiar and folded

upon itself ina way. If unravelled it would be longer than the

duodenal loop, and thus differs from that of Tachybaptes Auviatilis,

shown in the illustration (text-fig, 27). The extent of the ligament

uniting this with the duodenal loop is much as in Yachybaptes.

In the middle part of the intestine there are only two loops, as in

* Trans. Linn. Soc. fom. cié. p. 186.

+ Ibid. p. 185, fig. 5.

Proc. Zoo. Soc.—1911, No. VI. 6

82 MR. F. E. BEDDARD ON THE

Tachybaptes; Dy. Mitchell figures five. The two loops are wider

than those of Tachybaptes, but in the same way connected with

each other up to the veryend. Like Dr. Mitchell, I could find no

Meckel’s diverticulum.

Among the Ratu there is apparently but little modification of

the intestinal characters from genus to genus. Although I have

not examined the actual species which Hunter has described, it is

clear that his descriptions are in all probability perfectly correct.

Of the ‘‘ White Fulica [ Porphyrio albus, Cuv.|” he writes * : “The

duodenum passes down as usual, and then up, somewhat higher

than at beginning, making a sweep backwards to the loins and

commencing jejunum. This passes down on the right side, then up,

making a fold upon itself; then a second fold, as also a third,

which last is attached to the first fold: all these are parallel to

each other. The intestine then passes down, more in the middle

of the abdomen, further than the former three folds, along with

the duodenum.” This fully tallies in the number of loops with

Dr. Mitchell’s statement and with my own observations; these

latter are, I think, worth mentioning in brief, since they further

empbasise the uniformity of this group.

In Hydrornis alent, Rallus abbotti, Porphyrio melanonotus, and

P. madagascariensis the ileic and duodenal loops are pretty well

the same length and are attached by a duodeno-ileic ligament

nearly to the end of both loops. In all of these forms there are,

as both Hunter and Mitchell assert for species examined by

them, three loops only in the jejunal region of the gut. The

last of these three loops is the shortest of the three in all of

the species which I have just mentioned. In Porphyrio (both

species) I found a very conspicuous Meckel’s diverticulum, which,

as Mitchell correctly represents, lies near the bottom of the

middle loop. Finally, I may remark that this middle loop, at least

in Porphyrio, is free from the other two loops (as John Hunter has

stated), which are connected by mesentery.

Aramides ypecaha quite agrees with these other types and, as

in Porphyrio, the pancreas extends to the very end of the

duodenal loop. In this form also the first of the three jejunal

loops is the widest, and Meckel’s diverticulum occurs at about

the middle point of the middle one of the three loops.

Among the Gruss I have examined among others Anthropoides

paradisea and Balearica, which do not agree very closely in the

nature of their intestinal convolutions with those of Grus virgo,

as figured by Mitchell. In both these Cranes there are three, and

only three, jejunal loops, of which the first (as correctly indicated

by Mitchell) is much the widest. The two following are longer

and of equal length. On the first of these close to the blind end

of the loop there is (in Balearica) Meckel’s diverticulum. Mitchell

figures the ileic loop as trifid. I find that in both the Cranes and in

Grus japonicus which I have dissected (see text-fig. 18, p. 65) the

*® © Essays and Observations, p. 817.

i li

ALIMENTARY TRACY OF CERTAIN BIRDS. 85

ileic loop is a double loop, the two complete rather narrow loops

being not quite equisized. Anthropoides leacauchen agrees entirely

with the other two species, and in all of them the pancreas stops

some way in front of the end of the duodenal loop. The existence

of three separate loops in the jejunal region is obviously a point of

similarity between the Cranesand Rails and of difference between

the former and certain other ‘“Alectorides,” e.g. Cariama, Bustard.

SrpecanopopEs.— The intestinal tract of a species of Hregata has

been examined and reported upon by Dr. Mitchell, who, however,

was not in a position to state precisely to which species his

observations referred. J have examined an example of /regata

aquila, and have something to add to the facts enumerated by

Dr. Mitchell. He figures and refers to two ceca. I found only one

cecum, which was short and sessile, upon the gut. The duodenal

and ileic loops are simple and of about the same length. The

pancreas extends only about halfway down the duodenal loop

as in Birds of Prey, and the duodeno-ileic ligament is about co-

extensive with the pancreas. The duodenum is also connected

with the jejunal area by a ligament which I have not found in

other birds.

The condition of the jejunal area is interesting when compared

with that of other Dysporomorphous birds. It is, comparatively

speaking, short, while that of Cormorants and Pelicans is long.

This comparative shortness is mentioned by Mitchell, who, however,

has not seized upon a difference of some significance, as I think it,

which this bird shows from its allies.

In Phalacrocorax and Pelecanus (the only other genera ich I

have examined with reference to the matter now under con-

sideration) the jejunum is disposed in a considerable series of

closely applied regular fixed loops, as in Ducks, Storks, and some

other birds. Fvregata presents us with a stage anterior to this.

There are no definite and regular fixed loops, but the whole

jejunum can be disposed in an irregular circle with bulgings here

and there. It is not a simple archaic jejunum, as in the

Gallimaceous bird; but neither is it the much specialised jejunum

of other Dysporomorphe. It is hardly more advanced in the

direction of its immediate allies than is the corresponding part of

the intestine in an Eagle or an Owl.

The Psrrract possess a complicated intestinal tract, which is,

as I think, more correctly described by Owen than by Mitchell.

For it is quite impossible to represent the various loops into which

the intestine is drawn in these birds in the fashion adopted bv

Mitchell, as will be readily seen by a comparison of the

accompanying figure with his illustrations of Ara ararauna* and

Stringops ‘habroptilus ft. Owen mentions the ‘ packet of folds”

which are alternately connected as shown in the text-figure

appended, while Mitchell represents a series of loops sometimes

bifurcate or trifurcate, though stating that they are ‘“ folded

* P. Z. S. 1896, p. 155, fig. 21.

+ Trans. Linn. Soc., Zool, (2) viii. p. 244, fig. 61

84 MR. F. E. BEDDARD ON THE

upon each other, and twisted and doubled in a complicated

fashion.’”’ It is easy to state the characteristics of the Psittacine

alimentary tract, which in the species and genera Ara ararauna,

A. militaris, Nestor notabilis (see text-fig. 28), Chrysotis inornatus,

Psephotus hematonotus, Callocephalon galeatum, Platycercus

aveolus, Cacatua sulphurea, is constructed as follows.

The duodenal and ileic loops are single and elongated, as shown

in Mitchell’s figure. The jejunal loop consists of, proximally, a

series of three loops one within the other, of which the ascending

limbs of two are connected respectively with the duodenal and

ileic loops, and, more distally, of two separate single loops longer

than those of the proximal bunch, which are not directly con-

nected with each other but with the loops of the proximal packet.

The illustration will explain these interconnections better than a

more elaborate description,

Text-fig, 28.

oe seater

Intestinal tract of Nestor notubilis.

Lettering as before.

While there is in the species mentioned the general uniformity

of structure which has just been explained, there are differences

of small detail. Thus the two species of Aru differ, in that the

proximal complex of loops in 4. ararauna consists of only two

ALIMENTARY TRACT OF CERTAIN BIRDS. 85

short loops, from which it follows that the two long loops inter-

communicate directly. These facts, so far, are correctly shown in

Dr. Mitchell’s figure referred to. Conwrus leucotis is precisely

similar to Ara ararauna.

Melopsittacus undulatus is still further reduced. The proximal

complex is, as in the last species, reduced to two, but there is only

one of the two distal loops left and that is shortened.

On the other hand, Hclectus pectoralis is more complex than the

forms already considered. Among this group of birds the liga-

mentum ileo-duodenale is rather short and the pancreas extends

to the very end of the duodenal loop, or in some cases nearly so.

The ileic loop is often considerably longer than the duodenal. It

seemed to mé to be particularly long in Stringops habroptilus,

where it measured fully 8 inches in length.

Among the Piérons I have examined one or two forms not

studied by Dr. Mitchell, This group contrasts, for instance, with

the Rallide in the variety of intestinal patterns which it exhibits,

This, it will be noticed, is in accord with variations in the group

in other anatomical Chevacren *, I propose, however, to deal with

two genera in which the intestine has become shortened in relation

to the fruit-eating habit, leaving other forms aside until I have

been able to make a more comprehensive study. In one of these,

Ptilopus bellus, the reduction in length, coupled with great increase

in calibre, has not gone so far as in the genus Carpophaga, with

which I shall deal later. In Ptilopus bellus the duodenal loop is

quite well marked, though short and rather wide; the pancreas

extends to its very end. The jejunal region which follows is also

distinct from it and from the ensuing “ileic loop, The jejunal

region consists of a single loop only, the two limbs of which le in

close apposition, but can be considerably separated without tearing

any membranes. This region of the gut is therefore, in consisting

of a single loop without fir ther complications, pr ecisely like that

of the Bustards and Cariamas. The ileic loop is also well marked

and rather wide ; it is much longer than the duodenal. The ileo-

duodenal ligament is present, but not very extensive. There is

also a ligament uniting the jejunal loop to the entering limb of

the ileic “loop for about halfway down the latter. The plan of

the intestinal tract:in this bird is therefore a rather primitive one,

reminding us of that of Otis and Chunga and of the Picopasséres.

In fact, the degeneration of the intestine has resulted in the throw-

back to a pr esumably earlier state of affairs.

The genus Carpophaga shows a further degeneration of the

intestinal tract, which is well known to be very short in

this genus: of Pigeons ¥ r. The shortening is also accompanied by

widening.

I have examined C. enea and C. concinna: In both of these

the duodenal loop has vanished and is represented perhaps by the

* Garrod, “On some Points in the Anatomy of the Columbe,” P. Z. S. 1874, p. 249,

+ Cf, e.g., Garroi, “* Notes ov the Gizzard &c. of Car pophetga latrans,” PUzes.

1878, p. 102.

86 MR. F. E. BEDDARD ON THE

slightest kink in the alimentary tube. The rest of the tube is

disposed in four parallel and transversely arranged lines, which on

further examination are seen to form a spiral and which end in

a longitudinal section of gut running to the cloaca. I cannot

distinguish in this a jejunal from an ileic region. It is interesting

to compare with this “degeneration” of the intestinal tract in

the fruit-eating Pigeons the corresponding alterations met with

in the fruit-eating Passerine Zluredus. In that bird there is

a vestige left of the duodenal loop, to the extremity of which the

pancreas reaches. Indeed, the rest of the gut has retained,

though in a reduced condition, the two recognisable divisions, of

which the ileum is represented, as in the primitive Picopasseres,

by a straight tract parallel with and connected by the usual

membrane to the duodenal loop.

S$ Some General Considerations.

The new facts which have been described in the foregoing

pages give rise to certain reflections upon the affinities which

they appear to indicate between different families of birds and

upon the course pursued in the evolution of the gut of birds.

The facts, so far as they are known, do not appear to me to lead

to the establishment of a phylogenetic scheme, even of the

evolution of the gut only, so elaborate in the setting forth of

details as that which is drawn up by Dr. Mitchell as the result of

his own labours in this department of anatomy. Nevertheless, it

does seem possible to indicate certain stages in the evolution of

the intestine, and here and there are indications, already to some

extent considered, of affinities between different Avian families.

§ The Primitive Form of the Intestine in Birds.

It is obviously necessary, before considering the features which

are the most primitive in the Avian alimentary tract, to get a

clear notion of the essential differences which distinguish the

alimentary tract of Reptiles from that of Birds ; by this means we

shall evidently arrive at the essential resemblances. Dr. Mitchell

distinguishes the bird’s intestine thus :—-‘‘ It is distinguished from

the intestinal tract of reptiles chiefly by the fact that the three

divisions—the duodenum, Meckel’s tract. and the rectum—are

sharply marked off one from the other.” These lines are written

of Palamedea, which that author regards ‘‘as representing closely

the ancestral type.” It appears to me, however, that while

Palamedea is undoubtedly an ancient type, the definition used by

Dr. Mitchell is not a correct one. For, while in, at any rate, the

majority of Lizards known to me there is a very marked dis-

tinction between the small intestine and the large, the Crocodiles

show a further differentiation ; for they show a very well-marked

duodenal loop as well. The divisions of the alimentary tract

therefore do not enable us to distinguish between Birds and

ALIMENTARY TRACT OF CERTAIN BIRDS. 87

Reptiles. Indeed, in an earlier paper *, Dr. Mitchell has himself

sketched in a perfectly correct fashion the alimentary tract from

an Alligator, illustrating the facts to which I have just directed

attention. From a gut like this it appears to me to be only just

possible to distinguish that of Caswariust in its general charac-

teristics, among which I do not include the well-developed ceca.

The only difference that I can detect is a closer approximation

between the ileic and duodenal regions in Caswarius, already,

however, marked, though to a less extent, in the Crocodilide,

which, of course, foreshadows the very close association found in

all other birds. This association, caused by the outgrowth of the

long middle part of the small intestine from a short region of

the primitively straight gut, naturally brings about the com-

mencement of the formation of the fixed ileic loop, so conspicuous

a character of the alimentary tract of other birds. It is most

interesting to notice that among Crocodiles there is, in some

species at least, a quite distinct ileic loop, related perhaps to this

same association between the ileic and duodenal regions, which

is not, however, as has been already remarked, so close among

Birds. It is to be noted that here as elsewhere the closest asso-

ciation of Birds and Reptiles is shown, thoroughly justifying the

views of Cope, Huxley, and others. It may be admitted, therefore,

that Casuarius is, at any rate, one of those birds whose intestinal

tract, both arrangement and convolutions, hardly differs from

that of Reptiles, and is therefore primitive as compared with that

of many other birds. Nor, indeed, is there so far any very great

difference from the most primitive form of the gut in Mammals,

where, as in Casuarius and Crocodilus, the entire intestinal tract

is borne upon a continuous mesentery.

$ The Course of the Evolution of the Gut.

From the simple conditions which obtain in Cuswarius the

more complicated intestinal tract of other birds can be derived ;

and an almost complete chain of intermediate stages is exhibited,

even among the few genera which I have had the opportunity of

studying. It is from this point that the characters of the intes-

tinal tract in Birds diverge from those of Mammals, the Reptilian

conditions being left behind by both groups of Vertebrates. It

may be convenient at this stage to point out the essential

differences which distinguish the intestinal tract of Birds from

that of Mammals. It has already been pointed outt that one

difference is to be seen in the fact that among Mammals the

permanent loops of the large intestine distinguish that gut from

the small intestine, where there are no such permanent loops ;

whereas in Birds it is the small intestine only which ex-

hibits these permanent loops. These characters, however, though

* P.Z.S. 1896, p. 187, fig. 1.

+ I have examined the species C. vothschildi, C. intensus, and C. westermanni.

{ £.9., Gegenbaur, Vergl. Anat. d. Wirbelth.

88 MR. F. E. BEDDARD ON THE

distinctive in the negative sense are not universal. There are

whole orders of Mammals, e.g. Carnivora, Marsupials, Primates,

Edentata, in which the colon has no permanent loops, while in

many birds, ¢.g. Gallinaceous birds, certain Picopasseres, the

small intestine has no fixed loops. In addition to this very

important difference, there appear to me to be two other distin-

guishing features in the alimentary systems of the two orders,

which are perhaps equally important and are certainly more

regularly met wita. In all Mammals above those few forms

(e.g. certain carnivorous Marsupials, certain Edentata* ), which

have so far retained the Reptilian characters of gut, the whole

intestine is rotated upon itself and the rotation is fixed and the

change of position of the various sections of the gut retained by

the ligamentum cavo-duodenale which moors the end of the

duodenal loop to the colon, mesocolon, or median dorsal body-wall

on or near the postcaval vein. This ligament correlated with

the said rotation is universal among the higher and present in

most of the lower Mammals. In Birds, on the contrary, there is

no such tract of mesentery fixing the duodenum to the colon. So

far, in fact, the bird’s intestine has retained the primitive Reptilian

condition. The bird’s intestine, however, usually has what the

mammal’s intestine has not, a duodeno-ileie ligament. This doubt-

less is the physiological equivalent of the duodeno-colic ligament

(as Mitchell has pointed out), in so far as it serves to anchor the

perhaps otherwise inconveniently long and coiled small intestine.

It may also perhaps be argued from this that a short intestine

(e.g. many Picopasseres) is shown to be a secondary state of affairs,

from the very fact that it is in those birds provided with a duodeno-

ileic ligament, which may not be a mechanical necessity. For

the existence of the ligament in question may be due to mecha-

nical needs in an ancestor with a long small intestine. In any

ease, the morphological fact is to be noted and it constitutes a real

difference between the Mammalian and Avian gut.

In very nearly all birds whose intestinal tract is at or above

the level of that of the Cassowary, the end of the small intestine £

is attached by a mesentery of varying degree of development to

the duodenum. This, without any further specialisation, is the first

stage in the evolution of the gut from its simple archaie form.

This stage characterises the hypothetical Gallinaceous birds in

which it is universally present and quite similar in all.

This simple stage, which we may term Stage A, is also found in

other groups, but it does not occur in all the members of any

other group as it does in the case of the Gallinaceous birds.

Among the Struthious birds, for example, we have it in the

Cassowaries, Ostrich, and Apteryx§, whose intestinal tracts are

* Fora general survey, see Klaatsch, Morph. Jahrb. xviii. 1892, and myself in

P. Z.S. 1908, p. 568 &c. :

+ Trans. Z. 8S. xvil. p- 524.

+ The chief exception known to me is fuwmished by Fratercula arctica, the

remarkable characters of the intestine of which bird I have already commented

upon (supra, p. 78).

§ The condition of Rhee requires perhaps further study,

i ai alae,

ALIMENTARY TRACT OF CERTAIN BIRDS. 89

precisely similar so far to that of the Gallinaceous birds. We

have in certain Picopasseres (e. g., Melanerpes) the same state of

affairs, coupled in this case with a considerable shortening of the

eut. Among the Limicole the genus Plwvianus is also in this

stage.

The next stage, which may be called Stage B, is like the last,

save for the fact that the ileic loop is definitely formed. The

jejunal region remains unspecialised. To this stage, we refer the

gut in the majority of Picopasseres, including the Cuckoos and

Plantain-eaters. The ‘“Alectorides” (in the sense in which I venture

to use that term in the present communication) seem, but perhaps

only seem, to belong to this stage. Among the Limicolous birds

Sarciophorus and Gulls appear to belong here. Perhaps we should

also place in this assemblage the Dysporomorph /regata.

Stage C isa slight advance upon the foregoing. It is exem-

plified in certain Owls and Hawks, where the ileic loop is fully

differentiated off and attached in the usual way to the duodenum,

and where the jejunum is largely laxly coiled without any definite

loops, save a single loop, and that not a very well-marked one,

which occurs at the commencement of the jejunum. I have not

noticed this kind of intestine in any other group, except the

Nocturnal and Diurnal Birds of Prey.

Stage D might possibly be further divided up, but for the

present I do not see a clear way through the great variations

which the intestine of the more complicated forms shows. In all

of them the jejunum has become differentiated into fixed loops,

which vary in number, in relative length, and in their relations

one to the other. The majority of the larger birds belong to this

stage, as, for instance, the Cranes, Rails, Ducks, and Storks.

§ The Mutual Affinities of Avian Families judged by the

Intestinal Convolutions.

The known facts do not, as I think, permit of any complete

scheme of classification of Birds by means of the variations in

the coils of the intestinal tract. Here and there, however, there

would seem to be such indications, which are tolerably well

marked. More frequently, however, either a general plan runs

through a series of two or three groups, which makes any

definite placing of these groups in reference to each other

difficult, or a complete isolation is shown. The most salient

instance of the latter conclusion is undoubtedly the group of

Parrots, whose intestinal coils are constructed upon a plan which

is apparently universal in that group, but totally unlike anything

which is to be found in any other group. The affinities of the

Psittaci have been very variously interpreted *, but. it is clear

that the gut does not enable one to decide upon any of these

diverse views. J cannot at all agree with Dr. Mitchell in saying

* Many or most of these opinions are mentioned by Fiirbringer in_ his

monumental work, ‘ Uitersuchungen ueber Morph. u. Syst. der Végel,’ Amsterdam,

1888.

90 MR. F. E. BEDDARD ON THE

that “the relation to the common type is, however, easily made

out ” *,

In the same way, the Ralli are a quite circumscribed group

judged by their intestinal coils, which bear only a general resem-

blance to other groups and, indeed, to no group in particular.

Their characters are constant throughout the group, so far as

known facts enable us to make a statement. The most that can

be said is, perhaps, that they are nearer to the Grebes than the

latter are to any other subdivision of the Bird tribe. On the

other hand, it seems to me to be quite clear that if judged by

their intestinal coils the newer ideas with respect to the New

World Vultures and other Accipitrest must be abandoned, as

Dr. Mitchell has correctly pointed out; it is necessary to revert

to the older view which regarded the Condors as merely Vultures.

In the same way, although in this Dr. Mitchell does not agree

with me, the older opinion as to the Owls, that which placed

them close to the Accipitres and not in the neighbourhood of

various Picarian genera, is most certainly justified by the close

sunilarities in the mode of arrangement of the intestinal loops.

At the same time, it is also easy to distinguish these two groups

by the small but constant characters afforded by the ileo-duodenal

ligament.

And, again, it is by no means possible to distinguish by cha-

racters that carry any conviction the intestinal tract of a Grebe or

Tern from that of the Owls on the one hand or large Passerine

birds on the other; while the Gulls and Terns on their side

offer resemblances to what I venture to term ‘‘ the other” Limico-

line birds. Negative features are perhaps more salient in making

a brief survey like the present. Thus it is clear that the

Tinamous are quite unlike the Gallinaceous birds and that the

genus T'urnix is equally to be removed from that assemblage.

The Bustards and Cariama, moreover, show no. particular likeness

to the Cranes, though the first two seem to be closely allied to each

other, as [ have already pointed out. It is noteworthy that all the

four typest of Struthious birds differ from each other as much

as would seem to be possible in view of the undoubtedly primitive

characters of the gut in all of them, with the possible exception of

Rhea.

§ The Relationship between the Gut and the

Nature of the Food.

When we contrast the intestine of a Penguin with its enormous

series of closely adpressed straight loops and the intestine of a

Bustard with only three short intestinal loops, there would appear

to be a very marked difference between a fish-eater and an

omnivorous bird, and thus a close relationship between the form

and length of the gut and the nature of the food eaten by its

* P. Z.S. 1896, p. 155.

+ These opinions are so well known that I need not quote what would have to be

a long list of books and memoirs.

ft Casuarius and Dromeus belong, of course, to the same type.

ALIMENTARY TRACT OF CERTAIN BIRDS. 91

possessor. There is not, however, in all cases a relationship

which is plainly to be recognised between differences in diet and

differences in the intestinal part of the alimentary tract, and on

the other hand a totally different diet sometimes coincides with a

close similarity in the intestinal convolutions. Phylogenetic

relationship appears to me to have much more to do with these

similarities and differences in the gut. Furthermore, the way in

which the intestinal tract is modified in accordance with the diet,

when it does appear to be so modified, seems to have pursued a

different path in different groups in some cases. Of fish-eating

birds, for instance, the general idea is that the gut is long, and

that undoubtedly is the case with the Penguin and the Cormorant—

Pelican group. But then in these groups no representatives are

known which are not fish-eaters. Directly we come to the

consideration of groups of birds which contain fish-eating genera

and genera whose food is not fish, we are sometimes met by a

totally different state of affairs.

According to the statistics collected by Mr. Newstead*, the

Terns (of three species) are exclusively fish-eaters. Yet their gut

is not markedly and indeed hardly at all different from that

of the Avocet, which devours aquatic insects, and some other

Limicole which select a similar diet. The Kingfisher (Alcedo

ispida), which is, according to the same authority, practically

entirely a fish-eater, has a gut which is very like that of the

omnivorous Corvines, and has, moreover, a much shorter spiral

jejunum than in the differently feeding Halcyon sancta. The

Toucans ft are mainly fruit-eaters, though, like so many birds,

they will vary this diet with animal food. And yet their intes-

tinal tract differs very little from that of Podargus, which is

presumably not at all a fruit-eater, but subsists entirely upon

insects and other animals. Again, the Touracous? are fruit-

eating birds; but their gut is like that of a vast series of Pico-

passerine birds which feed upon all xin«s of food.

In asserting that the gut is short in all purely frugivorous and

insectivorous birds, Dr. Gadow practically admits how little stress

can be laid upon the relationship between length of gut and the

nature of the food. For the nature of the diet in each case is as

different as possible. Nor can any general principles be stated as

to the complication of the gut in tamilies of birds which live

differently. Thus the plan of the gut in Apteryx is practically

identical with that of the Gallinaceous birds, and the character

of the food differs. On the other hand, the pattern of the gut in

Accipitrine birds is not dissimilar to that of Owls, and here we

have a general similarity in diet. It is, in fact, not possible to lay

down general rules which have not copious exceptions. Many of

these exceptions can be gathered from the foregoing pages.

* Supplement to the Journal of the Board of Agriculture, vol. xv. No. 9 (1908).

+ Newton, ‘A Dictionary of Birds’ (London, 1898), sub voce * Toucan.”

£ Id. ibid., * Vouracou.”

» § Id. ibid., “ Digestive System.”

92 ON THE ALIMENTARY TRACT OF CERTAIN BIRDS.

§ Summary of Facts relating to the Intestinal

Coils of Birds.

We are in a position, I think, to lay down with confidence the

following general statements with regard to the intestinal tract

of the Class Aves. These statements are deduced from the

memoirs of others who have written upon this subject, as well

as from the facts ascertained by myself, and embody in a brief

form what is known upon the subject dealt with in the present

communication :—

(1) There are no essential differences between the intestinal

tract in Birds and in Crocodilia. The most complicated alimen-

tary tract in Birds can be derived through a series of stages from

the simple Crocodilian form.

(2) The intestinal tract of Birds differs from that of Mammals

in that there is never in the former, as there is generally in the

latter, a rotation of the gut coupled with an attachment of the

duodenum to the colon or mesocolon by a duodenal caval ligament.

On the other hand, there is nearly always in Birds an ileo-

duodenal ligament which serves to hold together the gut and

which is wanting in Mammals. Furthermore, in Mammals the

colon may be specialised into fixed loops, while in Birds such

fixed loops are found only in the small intestine.

(3) The small intestine of Birds can, with rare exceptions, be

distinguished into duodenal, jejunal, and ieic regions.

(4) The duodenal region consists of a long well-defined loop

(only absent in certain fruit-eating Pigeons), which is usually

straight, but sometimes (e. g. Milvus) shows indications of—or a

pronounced—spiral arrangement, as it does also in certain

Crocodilia. It is wider or narrower in different groups, and the

enclosed pancreas extends a greater or a less way towards the

extremity of the loop in different birds.

(5) The jejunal region is sometimes (¢. g. Gallinaceous birds)

marked off from the duodenal by a sudden diminution in calibre.

Tt is usually the longest section of the gut ; but is sometimes as

short or even shorter than either the duodenal or ileic region or

both. It may be a tract of intestine without any fixed loops (as

in the Gallinaceous birds, various Passerines, &c.), but if of con-

siderable length is usually specialised into a series of two or more

fixed narrow loops, which may be interconnected by secondary

mesenteries in different ways, distinctive of different groups of

birds. An intermediate condition is seen (e.g. among Accipitres

and in Fregata), where the jejunum is not arranged in lax and

alterable coils as in the Gallinaceous birds, but is stiffened here

and there into wide loops, which are not so sharply marked off

as in the more specialised birds and are not interconnected by

secondary mesenteries.

(6) The ileic region is not always quite distinct from the

jejunal, and degrees of distinctness occur. In less specialised

intestines there is no definite loop, but the end of the ileum is

ON SPOTTED HY ANAS IN THE BRITISH MUSEUM, 93

attached for a variable distance by a secondary mesentery to the

duodenal loop. In more specialised intestines the ileic region

forms a distinct loop which is narrower or wider, and which is

equal to, longer, or shorter than the duodenal loop, to which it is

attached for a greater or less length by the ileo-duodenal ligamené

already mentioned. Not infrequently the ileic region consists of

two loops, of which one is often small and lies just above the

creca.

(7) The plan of the gut is constant (except for very slight

variations) in all the members of certain groups. ‘This isthe case

with the Psittaci, Galli, Raptores, Striges. In the case of other

groups, there are considerable divergences of structure within the

group: this is the case with the Picopasseres, Limicole, Grues,

Struthiones (if, that is to say, the two latter groups can be

regarded as natural groups, which is open to doubt on other

grounds).

(8) A comparison of the intestine of Birds with that of

Reptilia (especially Crocodilia) allows of the recognition of more

and of less primitive types of intestine. The most primitive type

is found in the Cassowaries, Struthio, Apteryx, and all the

Gallinaceous birds; and is also seen among the Picopasseres

(Woodpeckers) and Limicole (Pluvianus). Most Picopasseres

and the Birds of Prey (nocturnal as well as diurnal) show a rather

more specialised form of intestine. In the remaining groups of

Birds the intestine is more specialised still and in several different

directions.

(9) Certain classificatory results seem to follow from a

comparison of the differences exhibited by the intestinal tract.

Thus, the resemblance of both Cuculi and Musophagi to the

Picopasseres, and the likeness between all the Accipitres (New

World and Old World, nocturnal and diurnal) are remarkable.

The close likeness between the Bustards and the Cariamide is

to be commented upon. The Passerine character of the gut of

Turnix and the possible likeness between Crypturus and Rhea

seem also to be shown.

7. On the Specimens of Spotted Hyzenas in the British

Museum (Natural History). By Prof. ANGEL Caprera,

C.M.Z.S.

[Received November 5, 1910: Read November 29, 1910. |

Every zoologist working on the Spotted Hyzenas with suitable

material from different localities feels the convenience of re-

cognising several local forms. As early as in 1812, two of them

were admitted by Cuvier, and in modern times no less than nine

other “species” have been described. It is not easy to say with

which form Erxleben’s Hyena crocuta, afterwards the type of the

genus, or subgenus, Crocuta, must be identified, as the species was

94 PROF. A. CABRERA ON

based on the “ Spotted Hyena” of Pennant*, and this author

gave no definite locality, saying only that the animal is found in

“ Guinea, Althiopia, and the Cape.”, That indication practically

embraces all the African countries known in Pennant’s time,

excepting only Barbary and Egypt. As to the original de-

scription, made from a specimen shown in London some years

before, it runs thus :—

“Short black mane: hair on the body short and smooth: ears

short and a little pointed; their outside black, inside cinereous :

face, and upper part of the head, black: body and limbs reddish

brown, marked with distinct round black spots; the hind legs

with transverse black bars; tail short, black, and full of hair.”

Now, I have never seen, nor found described, a Spotted Hyzena

with black mane. Young specimens commonly have dark hairs

in it, producing a general blackish tinge, but it seems clear that

Pennant’s specimen was not young, as in the description it is

afterwards stated that it was bigger than the striped species, and

the author says about the latter in a previous page that it 1s

larger than a big dog. It is therefore necessary to suppose either

that Pennant spoke from memory and forgot some details of the

coloration, or that the actual specimen represented a form quite

unknown to modern naturalists. The latter view being a very

unlikely one, I prefer to think that the description was written

from memory only, under the impression of a reddish-brown

animal spotted with black, and perhaps a not quite developed

specimen with a little of the juvenile dark hair in the mane.

This determination being adopted, it seems to me very probable

that the specimen alluded to came from Senegambia, although

this locality is not mentioned among those given by Pennant as

inhabited by Spotted Hyznas. Senegambia and the Cape were,

during the eighteenth century, the two countries that chiefly and

almost exclusively furnished the European menageries with

African animals. But in the Cape Hyena the ground-colour

is a dirty yellowish which nobody would call reddish brown,

whereas this rather indefinite designation may be correctly

applied to the peculiar colour, intermediate between dark

cinnamon and raw umber, of the Senegambian Hyzena. ‘The fact

that Pennant did not include Senegambia in the habitat of the

species is of little, if any, importance, as he compiled the geo-

graphical distribution from the works of Bosman, Kolbe, &c., and

was evidently unaware of the provenance of the specimen he saw

in London. In his ‘Game Animals of Africa,’ Mr. Lydekker says

that the typical Hyena crocuta is the form found from Southern

Egypt, across Central Africa, to Senegal in the west and the

Transvaal in the south. I cannot agree entirely with such a

conclusion. As will be seen below, Spotted Hyznas from the

Nile Basin and East Central Africa are very different in colour

from the animal described by Pennant and Erxleben, and therefore

SPOTTED HYNAS IN THE BRITISH MUSEUM. 95

these countries at least must be discarded. The supposed dis-

tribution being restricted in this manner to Central Africa and

Senegambia, it becomes clear that, as Central Africa was totally

unknown to Europeans in Pennant’s time, the only conclusion at

which we can arrive from Lydekker’s statement is that which I

have adopted. Crocuta crocuta may therefore be selected as the

name for the Senegambian Spotted Hyena, at least until a

stronger argument can prove that this view is not the right one.

As I have said above, Cuvier, in his ‘Ossements Fossiles,’

distinguished two different forms of Spotted Hyznas, a grey

one and a reddish one. In the second edition of the same work,

he states that his reddish Hyzna is frequently found about

the Cape, but the source of this knowledge is not given. The

first exact description of the Cape Hyena is by Desmarest*, who

named it Hyena capensis. As to H. rufa, of the same author,

based on Cuvier’s reddish Hyena, its exact locality being unknown,

it is best to consider it as a synonym of true erocuta. Boitard +

gives the three names to one and the same animal, and describes

a yellowish Hyena from the Cape as H. cwvieri, this name

becoming thus a synonym of Desmarest’s capensis.

Since the publication of all these old names, no other splitting

of the group has been attempted till 1900, when Matschie = de-

scribed five so-called new species: Crocuta wissmanni, from German

West Africa; C. gariepensis, from the Orange River; C. ger-

minans, from German Hast Africa; and C. thierryi and

C. togoensis, from Togo. In a subsequent paper §, the same

zoologist named the form from Kamerun C. noltei. Satunin || has

given the name leontiewi to the Abyssinian Spotted Hyena, and

Lonnberg 4 has described two other forms from East Africa,

C. kibonotensis, from Kibonoto plains, and C. panganensis, from

the Pangani River.

It is impossible to decide now, without the comparative study

of large series of specimens, and especially of skulls, whether all

these forms are true different species, or whether they are local

races of one or of several species. From the material in the British

Museum, which I have been able to examine through the kindness

of Mr. Oldfield Thomas, F.R.S., it appears that two different types

of skulls may be distinguished :—a broad one, in which the width

of the palate across the upper carnassials is practically equal to or.

a little greater than the length of the lower tooth-series exclusive

of incisors ; and a narrow one, in which the width of the palate is

equal to or a little less than the length of the upper tooth-series.

This latter always being 4-15 mm. shorter than the lower tooth-

row, it is clear that the difference between the two types can be

* “Mammalogie,’ i. (1820) p. 216.

+ Le Jardin des Plantes, 1845, p. 232.

{ SB. Gesellsch. Nat. Fr. Berl. 1900, pp. 18-58.

§ L. ec. 1900, p. 211.

* Zoologischer Anzeiger,’ xxix. (1905) p. 556.

"| Sjostedt, Kilimanj. Meru Exped. 1908, pp. 16-18, pls. 5 & 7.

96 PROF, A. CABRERA ON

detected at first glance. Moreover, the ratio between the zygo-

matic breadth and the condylo-basal length is 65 to 72:5: 100 in

narrow skulls, and 70 to 75:100 in broad skulls. Of course, I

refer solely to adult specimens; in young skulls the proportions

are very changeable, according to the age.

Three only among the eleven forms described up till now

seem to be represented in the British Museu collection, as

follow :—

CROCUTA CAPENSIS Desm.

Two adult specimens, one from the Cape (B.M. no. 46.8.3.3,

Turner), and another from the Pongola River, Zululand (B.M.

no. 2.2.8.1, D. Bruce), and a young one from the Cape (B.M.

no. 37.9.26.90, Turner). Ground-colour diyty cream-buff ; spots

small, round, numerous, and of a pale hair-brown; mane dirty

ochre-yellow; snout and feet dark-coloured, between sepia and

hair-brown. Skull of the broad type.

Skull-measurements of no, 46.8.3.3.—Condylo-basal Jength,

236 mm.; zygomatic breadth, 176; interorbital breadth, 57 ;

postorbital constriction, 42; rostral breadth on canines, 64; width

of palate across the carnassials, 114; mandible, from condyle,

180; upper tooth-series*, 100; lower tooth-series, 111; upper

carnassial, 35 x 22; lower carnassial, 30 x 12.

A specimen from the Cape in the Madrid Museum of Natural

Science is entirely like the British Museum specimens. The

figure given by F. Cuvier in his ‘ Histoire Naturelle des Mammi-

féres,’ after a living animal obtained in the same locality by

Capt. Baudin, is not good, the difference in height between the

fore and the hind quarters being too exaggerated.

CROCUTA WISSMANNI Matsch.

An old specimen from Linyanti (B.M. no. 0.10.3.1, P. C. Reid).

Ground-colour ochraceous; under surface, from the throat, pale

cream-colour. The spots are very dark brown and show a

tendency to form longitudinal rows on the flanks. On the limbs

they are darker, almost black, and reach a lower level than usual

in the group. Mane ochraceous. Snout dark brown. Feet buff-

yellow. Skull of the narrow type.

Skull-measurements.—Condylo-basal length, 256 mm.; zygo-

matic breadth, 185; interorbital breadth, 64; postorbital

constriction, 53 ; rostral breadth on canines, 66; width of palate

across carnassials, 112; mandible, from condyle, 190 ; upper tooth-

series, 113; lower tooth-series, 117; upper carnassial, 35 x 20 ;

lower carnassial, 32 x 11.

It is not without hesitation that I call this specimen wissmanni,

as it lacks the black feet which Matschie says are characteristic

of the German West Africa Hyzena, and which are to be seen in

* In describing Carnivora, I always measure the tooth-rows from the front of

the canine to the back part of the hindmost cheek-tooth.

SPOTTED HY/ENAS IN THE BRITISH MUSEUM, 97

Schreber’s plate xcvi. (Saugth.), with which wissmann/ is identified

But the ochraceous ground-colour, the paleness of the under parts,

and, moreover, the relative proximity of Linyantito Epukiro, the

type-locality of wissmanni, prevent me from giving it a new name,

at least until some other specimens are available. By the way,

the material on which Crocuta wissmanni was based is far from

good. The type is askin without skull, and Prof. Matschie is

not sure that the skull from Windhoek, described in his paper,

really belongs to the same form.

CROCUTA LEONTIEWI Satunin.

A skull, without skin, from Abyssinia (BLM. no, 69.2.2.13,

Jesse). ts dimensions answer rather well to those given by

Satunin for his specimen 5784. It belongs to the narrow type

and is broken behind, it being, therefore, impossible to measure

its condylo-basal length.

Skull-measurements.—ZLy gomatic breadth, 153 min.; interorbital

breadth, 55:5; postorbital constriction, 41; rostral breadth on

canines, 58; width of palate across the carnassials, 99 ; mandible,

from condyle, 172; upper tooth-series, 98 ; lower tooth-series, 103 ;

upper carnassial, 36 x 19; lower carnassial, 26 x 10.

Another skull, without skin, from the White Nile (B.M. no.

2.8.5.4, Maj. Duni), seems to belong to the same species ; whereas

another from Bar-el-Zaraf (B.M. no. 0.8.6.2, Stanley Hlower)

evidently represents a different form, perhaps the Sudanese

Hyena, which I donot dare to name without knowing the colours.

It is of the broad type, and undoubtedly came from a very big

animal,

There are in the British Museum, besides the specimens just

mentioned, some others representing three different and ap-

parently new forms. In describing them, it is only provisionally

that I use binomial names.

CROCUTA RUFOPICTA, sp. n.

Diagnosis.—A very pale reddish form with red spots, quite

different from the grey Hyzenas of Abyssinia and Uganda.

Colowr.—Ground-colour pale ochraceous buff, verging to tawny

ochraceous on the back and the mane, and fading to pale buff on

the under parts. Spots small, very scattered, and ochraceous

rufous in colour. Feet ochraceous. Tip of the tail blackish, as

usual in Spotted Hyzenas.

Skull.—Palate narrow, its greatest breadth being rather less

than the length of the upper tooth-series. Bulle elongate ;

paroccipital processes considerably drawn backwards, their hind

outlines appearing, when the skull is resting on its basis, as

oblique lines.

Skull-measurements (of type).—Condylo-basal length, 240 mm. ;

zygomatic breadth, 159; interorbital breadth, 53:5; postorbital

constriction, 40; rostral breadth, 59; width of palate across the

carnassials, 97; mandible, from condyle, 178; upper tooth-series,

Proc. Zoou, Soc.—1911, No. VII, 7

98 ; PROF. A. CABRERA ON

103; lower tooth-series, 106°5; upper carnassial, 36 x 20; lower

earnassial, 28 x 11.

Hab. Odueina, Boran Country.

Type. Old female. B.M. no. 9.6.1.14. Collected by Mr. Drake-

Brockman.

Remarks.—The type-skin lacks the head and fore part of the

neck, but it is so different from all the other Spotted Hyzenas

that, although the only specimen and an incomplete one, it

deserves to be considered a new form. C. kibonotensis, pale

reddish in colour like rufopicta, has the spots dark brown, not red,

and its skull, an excellent photograph of which has been published

by Loénnberg, is also different from that of the present species or

race.

CRrocuUTA THOMASI, sp. n.

Diagnosis.—A pale grey, black-spotted Hyena, with the skull

of the narrow type.

Colowr.—General colour pale grey, strongly suffused with buff

on the fore quarters, and marked with large black spots, very

irregular in form, some of them being elongate and even some-

what linear. Mane dirty ochre-yellow, fading to cream-buff on

the sides of the neck, where there are two irregular rows of faint

burnt-umber spots. Feet and legs to near the elbow and the

knee dark clove-brown, almost black. Tail pale grey, blotched

with small blackish spots, and with the distal half black.

Skull.—Palate narrow, but not so much as in C. rufopicta, its

largest breadth being practically equal to the length of the upper

tooth-row, and even considerably exceeding it in young specimens,

in which the carnassial is not yet quite developed. Hinder out-

line of paroccipital processes forming a vertical line.

Skull-measurements (of type).—Condylo-basal length, 245 mm. ;

zygomatic breadth, 165 ; mterorbital breadth, 54; postorbital con-

striction, 41; rostral breadth on canines, 58; width of palate

across the carnassials, 101 ; mandible, from condyle, 175; upper

tooth-series, 100; lower tooth-series, 107; upper carnassial,

35x18; lower carnassial, 26 x 10°5.

Hab. Ankole, Uganda.

Type. Adult male, B.M. no. 1.8.9.27. Collected by Sir Harry

Johnston.

Remarks.—I have seen two other specimens, from Ankole also,

one obtained by Sir Harry Johnston (B.M. no, 1.8.9.28) and the

other, a young female, collected by Mr. Delmé Radcliffe (B.M.

no. 5.4.3.4). All three are readily distinguishable from the

other grey Hyenas of Hast Africa. The Abyssinian C. leontiewi

is not pale grey in colour, but “ bra’unlichgrau, auf der Mitte des

Riickens rostbraunlich,” and has a smaller skull (zygomatic

breadth about 155 mm., against about 165 in C. thomasi). In

C. panganensis, from the Coast Region, the colour is darker,

brownish ash, verging to rusty brown in the mane, and the skull

is considerably broader behind the postorbital processes. The

SPOTTED HY ZNAS IN THE BRITISH MUSEUM. oo

West-African C’. togoensis appears to be the only species similar

in colour to the Uganda Hyena, but according to plate 104 of

Dr. Heck’s ‘Lebende Bilder aus dem Reiche der Tiere,’ in

which the type of the species is figured, in the Togo Hyzena the

large spots are more regular in form, and there are among them

some very small spots which are not present in thomasi. The

skull of fogoensis is, moreover, of the broad, not of the narrow,

type. The original specimen is a young one, and therefore use-

less for comparison, but an old topotypical specimen in the Berlin

Museum, Prof. Matschie kindly informs me, presents the follow-

ing dimensions: width of palate across the carnassials, 107-1 mm. ;

upper tooth-series, 99°8; lower tooth-series, 109°3. Of course, it

was not to be expected that a so plastic group would be re-

presented by the same form in two countries so widely separated.

I have much pleasure in naming the Uganda Hyena in honour

of Mr. Thomas, as a remembrance of the kindness shown me while

examining the invaluable collections under his charge.

CROCUTA NYAS, sp. n.

Diagnosis.—A pale yellowish Hyzena, with large dark spots and

pale brown feet. |

Colour.—General colour dirty buff; mane pale ochraceous ;

belly dark brown. The spots are dark Vandyke-brown, large and

very scattered on the body, smaller and more close-set on the

limbs. On the sides of the neck there are some faint traces of

brownish-red spots. The muzzle is of a pale, dirty wood-brown,

and the feet are of this same colour, somewhat lighter.

Skull.—Palate long and narrow, its greatest breadth practically

equal to the length of the upper tooth-row. Zygomatic arches

comparatively close to the skull.

Skull-measurements (of adult female, paratype).—Condylo-basal

length, 260 mm.; zygomatic breadth, 179; interorbital breadth,

61; postorbital constriction, 48; rostral breadth on canines, 69 ;

width of palate across the carnassials, 110; mandible, from

condyle, 190; upper tooth-series, 111; lower tooth-series, 119;

upper carnassial, 38 x 20; lower carnassial, 32 x 12.

Hab. Mount Milanji, South Nyasaland.

Type. Immature skin and skull. B.M. no. 92.8.1.5. Col-

lected by Alexander Whyte and presented by Sir Harry Johnston.

Remarks.—There are in the British Museum, besides the type,

an old female skull (92.8.1.4), measured above, and a stuffed

specimen exhibited in the public galleries, both received from

Sir Harry Johnston. The species, which very probably is the

same as found in Portuguese Hast Africa, is readily distinguish-

able from C. germinans, its geographically nearest ally, this latter

being a much darker animal, wood-brown with the mane clay-

colour.

100 ON A VARIETY OF PLUMAGE IN A RED GROUSE.

EXHIBITIONS AND NOTICHES.

December 13, 1910.

G. A. Boutenerr, Esq., F.R.S., Vice-President,

in the Chai.

THE Seorerary read the following report on the additions made

to the Societys Menagerie during the month of November,

1910 :—

The number of registered additions to the Society’s Menagerie

during the month of November last was 181. Of these 72 were

acquired by presentation, 31 by purchase, 33 were received on

deposit, 35 in exchange, and 10 were born in the Gardens,

The number of departures during the same period, by deaths

and removals, was 209.

Amongst the additions special attention may be directed to :—

1 American Bison (Bison americanus), born in the Menagerie

on Nov. Ist.

5 Canadian Wapiti (Cervus canadensis), 1 g, 4 2, from Canada,

purchased on Noy. 8th. f

1 Tasmanian Wolf (Zhylacinus cynocephalus), ¢, and 2 Tas-

manian Devils (Sarcophilus satanicus), from Tasmania, purchased

on Noy. 21st.

Dr. H. Hammonn Surry, M.R.C.S., F.Z.S., exhibited a mounted

specimen of the Red Grouse (Lagopus scoticews) which displayed

a curious variety of the ordinary plumage, and made the following

remarks :—

“This Grouse was sent to me by Mr. Wynyard Dixon, of

Sheffield, from the Gilkerscleugh Moors, Abington, Lanarkshire,

on October 4th this year. The bird is a cock. The colouring is

not common, and the specimen is not quite like the so-called

pepper-and-salt variety, of which I saw a good example at

Mr. Boyals, where this bird was set up. In the bird now

exhibited, while the wings show grey colour, the feathers under

the chin are more like the colour of the feathers of a young cock

pheasant, and there is just an appearance of a light ring round

one side of the neck; there are also a few bronze-coloured

feathers on the flanks. Of this moor Mr. Dixon says in one of

his letters: ‘I was much surprised the first time I came to this

district to find pheasants on the moors considerable distances

from coverts or farms—ain fact, the pheasants do not appear to

come into the spinneys till November’; he further states that

he has seen them two miles from any covert. But all who have

shot on moors adjoining pheasant preserves are aware that

pheasants will stray for very long distances over the moors from

the coverts, especially if there are bilberries to be found. It has

been suggested that this bird may be a hybrid between the

pheasant and the grouse; J can find no previous record of such

eee eee

THE OCCIPITAL REGION OF THE HEAD IN BATRACHIA URODELA. 101

a hybrid. Grouse vary considerably in their colouring from very

dark to cream-colour. I do not personally contend that this bird

is a hybrid; in my opinion it is a Grouse, and I show it this

evening as a curious variety of the colouring of the ordinary

Red Grouse.”

Other zoologists present confirmed Dr. Hammond Smith’s

opinion that the bird was not a hybrid.

Mr. D. Sera-Surru, F.Z.8., Curator of Birds, exhibited some

skins of the Australian Yellow-rumped Finch (Jhunia flavi-

prymna). These birds had been kept alive in an outdoor aviary

in England, and had developed certain markings tending towards

those of another closely allied species, Munia castaneithorax.

The exhibitor attributed this to the fact that the former species

was a desert form of the latter, and when placed in a humid

environment tended to revert to the plumage of the latter. He

referred to a paper he had published on this subject in the

‘ Avicultural Magazine,’ 1907, p. 195.

Dr. W. E. Hovis, M.A., F.Z.8., English Member of the Inter-

national Commission on Zoological Nomenclature, explained the

Report presented to the Graz Meeting of the International

Zoological Congress, and referred in particular to the proposals

made for the protection of well known zoological names.

A discussion followed on the portion relating to the formation

of an Official List of most frequently used Zoological Names. The

feeling of the Meeting was very strongly in favour of the Inter-

national Congress giving its authority to the forming of a List of

Zoological Nees ane Sen tfcsnn of which should not be altered

by application of the rules of the International Code. It was

unanimously agreed to accept the action of the Congress if it

would adopt this course.

PAPERS.

8. On the Segmentation of the Occipital Region of the Head

in the Batrachia Urodela. By Hpwin 8. Goopricy,

M.A., F.R.S., F.Z.S., Fellow of Merton College, Oxford.

[Received November 29,1910: Read December 13, 1910. |

(Text-figures 29-51.)

Lntroduction.

It is now well known that in the Craniata Gnathostomata the

region of the head lying behind the auditory capsule is a

compound structure, formed of a number of segments originally

102 MR. E. S. GOODRICH ON THE OCCIPITAL REGION

like those of the trunk. A process of cephalisation leads to the

fusion and partial suppression of a number of skeletal segments,

or scleromeres, which combine into a compact occipital region

continuous with the remainder of the skull in front. Through

its wall issue segmental nerves. At the same time, there is a

tendency for the corresponding muscular segments to become

reduced. The history of this subject has been so often told that

it need not be repeated here (Sewertzoff 9, Gaupp 8, and

myself 6).

The occipital region in the Amniota has been found to include

behind the vagus nerve four scleromeres enclosing three roots of

the hypoglossus nerve. There are therefore probably at least

five segments altogether between the auditory capsule and the

atlas—the first corresponding to the glossopharyngeal, the next

to the vagus, and the last three to the hypoglossal. Possibly

there are a few more.

In the Pisces the posterior limit of the head is both less

definite and more variable in position; but the postauditory

region probably always includes at least seven segments. The

Selachians have been most thoroughly studied, and in them there

are about eight segments behind the auditory capsule (metaotic

segments). The first corresponds to the glossopharyngeal nerve

and the fourth mesoblastic somite (three of these being prodtic) ;

the next four segments belong to the vagus, and the last three to

the hypoglossus, much as in Amniotes. But in the Selachian the

anterior sclerotomes and myotomes are more distinct. The first

metaotic somite produces no myotome, and therefore preserves no

ventral nerve-root. The remaining occipital somites contribute to

the hypoglossal musculature, and their corresponding ventral roots

are the spino-occipitals of Fiirbinger (s—z). But the muscles of

these segments tend to disappear in ontogeny from before back-

wards. In adult Selachians some two or three hypoglossal roots

are usually found piercing the occipital region of the skull. An

examination of the early stages, however, reveals the complete

series of somites, nerves, and skeletal segments, which make

up the postauditory region of the head.

Very different is the state of things in the Batrachia (Am-

phibia). Here the skull appears to end immediately behind the

vagus foramen ; and nerves contributing to the hypoglossal issue

from the vertebral column behind the occipital condyles.

The first, second, and third spinal nerves may form the

complex hypoglossal; the second being the main, and often the

only, hypoglossal nerve in the adult.

These facts immediately suggest several questions :—Does the

occipital region of the Amphibian really include fewer segments

than that of the other Gnathostomes, or have certain segments

been telescoped and practically crushed out? Are the hypo-

glossal segments of the Gnathostomes really represented by the

first three trunk-segments of the Amphibian, or have these simply

assumed the function originally fulfilled by others farther forward ?

OF THE HEAD IN THE BATRACHIA URODELA. 103

Further, if the Amphibian head ineludes fewer segments, it may

be asked whether this condition is primary, or due to the return

of segments to the trunk which formerly held a place in the head.

It is essential before attempting to answer these questions to

determine how many head-segments can actually be traced in

ontogeny. Other observers have attacked the problem, but

their results are not in agreement. With a view to settling this

point I undertook the study of the development of the head-

region in the Axolotl (Amblystoma tigrinwm).

My method has been to reconstruct series of sections on

paper. For this purpose it is important to have a very com-

plete set of stages cut in various directions. I have to thank

Dr. J. W. Jenkinson for the loan of a large number of excellent

series of sections filling up the gaps in my own preparations. An

appropriate number of stages has. been selected for represen-

tation in the figures given in the text; but it will be understood

that intermediate stages have been examined. Of these figures,

nos. 33-38, 40-43, and 45-49 were drawn from Mr. Jenkinson’s

series.

Description.

Before describing my own observations it will be well to give

a brief account of the results obtained by previous workers. A

good general description of the development of the skull of the

Axolotl has been given by Parker (7). He describes the basilar

plate, or floor of the cranium behind the pituitary fossa, as formed

of a parachordal extension of the trabecular bars, combined with

“ proper occipital parachordals behind.” The latter give rise to

the occipital condyles ; but their exact relation to the myotomes

is not elucidated, although Parker notices that the glossopharyn-

geal ganglion lies behind the auditory capsule between the first

and second myotomes. Stdhr (11) has given a very similar account

of the development of the skull of Siredon (Amblystoma). Like

Parker he found that the postpituitary region of the skull develops

from three separate sources: the anterior parachordals (“‘ Balken-

platten ”) derived from the trabecule, the auditory capsules, and

the occipital arches. By means of reconstructions of sections he

made out clearly the origin of the posterior occipital arch. These

arches, compared to vertebral arches, grow over the brain above,

and along the sides of the notochord (occipital parachordals of

Parker) join the backward extensions of the trabecule (‘‘ Balken-

platten ”’), and enveloping the notochord form the basilar plate,

which subsequently fuses with the auditory capsules. Stohr,

however, did not make out the exact position of the occipital

element with regard to the nerves and myotomes.

Sewertzoff (9) was the first author to attack this problem directly.

According to his account there are two metaotic somites, giving

_ rise to two myotomes in the embryo (text-fig. 50, B, p. 116).

The Ist disappears later. The 2nd, corresponding to the vagus,

remains. Behind it, in the septum between the 2nd and

3rd, develops the occipital arch. The first trunk-segment

104 MR. E. §. GOODRICH ON THE OCCIPITAL REGION

(3rd metaotic) has a myotome, a ventral root, but no ganglion.

The next and succeeding trunk-segments are complete. That

this description is incorrect has already been suggested by Miss

Platt, who has given us a detailed and admirable account of the

development of the head in Wectwrus (8).

Miss Platt analyses the postauditory region as follows :—The

Ist somite belongs to the glossopharyngeal segment, develops no

myotome and has no ventral root. The 2nd, 3rd, and 4th are

vagus segments; the 2nd somite disappears ventrally, but its

dorsal portion develops muscle which combines with the more

fully formed myotome of the 3rd somite. The 4th and succeeding

somites develop myotomes. ‘The 3rd, 4th, and 5th grow down

ventrally to give rise to the hypoglossal muscles, supplied by the

ventral roots of the 4th and 5th segments. The 6th segment

(3rd of the trunk) is the first to have a complete spinal nerve.

with ganglion, ventral and dorsal root. The first neural arch hes

between the 4th and 5th somites. Between the 4th and 3rd

somites appears the occipital arch marking the hind limit of the

skull; while between the 3rd and 2nd somites is formed a rudi-

mentary preoccipital arch, which is taken into the auditory

capsule. If this account is correct, there are three metaotic

segments in the head of Amphibia, the myotomes of the last two

being represented in the adult by the anterior region of the dorsal

temporal muscle.

Text-fig. 29.

ine, aya Mh eHile

pal.

Dissection of a full-grown Axolotl, showing the skeleton and nerves of the head

and three trunk-segments. The cartilage is dotted. View from left side.

Coming now to my own observations on Amblystoma, we may

begin with a glance at the structure of the full-grown animal as

shown in text-figs. 29 and 30*. The hyomandibular branch of the

facial nerve issues from behind the otic process of the quadrate

* For explanation of the lettering of these text-figures see p. 120.

_—_— ee

OF THE HEAD IN THE BATRACHIA URODELA. 105

and the overlying squamosal, and passes down the hyoid arch. The

glossopharyngeal and vagus come out together behind the auditory

capsule—the former supplying the first branchial arch, and the

latter giving off three branches to the remaining arches. A large

visceral branch of the vagus runs along the alimentary canal, a

Text-fig. 30.

Outline of spinal cord and hind brain, with roots of cranial nerves and first two

spinal nerves, seen from left side.

dorsal branch upwards, and slender lateral line branches to the

skin. Through the bony neural arch of the first vertebra issues

the first spinal nerve, having a ventral root only. Passing over

the vagus it soon joins the complete second spinal, which has

two roots *, and comes out behind the first vertebra. A. dorsal

twig of the first spimal supplies the temporal muscle. The first

and second spinals together make up the hypoglossal nerve.

Text-fig. 31.

Partial reconstruction of the hind region of the skull and the anterior region of

the trunk of an advanced larva with a head 6 mm. long. The cartilaginous

skeleton and nerves are shown through the myotomes.

Text-figs. 31 and 32 are partial reconstructions of a larva which

has nearly acquired the structure of the fully grown animal.

The auditory capsule is chondrified and fused to the basilar plate,

* Miss Platt finds only a ventral root to this nervein Necturvs. Theré can be no

doubt that both roots are present 11 Amblystoma as well as a eanglion.

106 MR. E, S. GOODRICH ON THE OCCIPITAL REGION

Text-fig. 32. .

Partial reconstruction of the hind region of the skull and the anterior region of the

trunk of the advanced larva shown in text-fig. 31. The myotomes have been

removed.

Text-fig. 33.

Sg)

mt 8? mt.

i !

]

” Git ac,

Partial reconstructions of the left side of the head region of a younger larva,

in which a large amount of cartilage is developed.

Text-fic. 33 shows the inner wall of the auditory capsule, and the anterior myotomes

of the trunk through which are seen portions of the skeleton and spinal

nerves. The vagus nerves have been cut short at their exit from the skull.

In text-fig. 34 the skeleton and nerves are more completely shown, but the

muscles have been removed,

OF THE HEAD IN THE BATRACHIA URODELA. 107

and the preoccipital and occipital arches. The latter is complete

above the brain. The second muscular segment, really the third

myotome developed in the 4th somite, is split in two by the

vagus. The first hypoglossal nerve comes out between the skull

and the first neural arch.

esto ae

ne poc. gt. na. nch,

Partial reconstruction of the left side of the head region of the larva shown in

text-fig. 34. Represents the same structures cut back to nearer the middle

line, the auditory capsule and side wall of the skull being removed and the

vagus roots exposed,

Text-fig. 36.

Reconstruction, seen from above, of the hind region of the skull, and anterior

neural arches, Some roots of cranial nerves are seen on the right.

108 MR. E. 8S. GOODRICH ON THE OCCIPITAL REGION

A larva slightly younger is shown in text-figs. 53, 34, and 35.

At this stage the ventral portion of the 2nd myotome is still

preserved. It les below the vagus between the occipital and the

preoccipital arches. ‘The latter is seen to be a thickening in the

inner wall of the auditory capsule, continuous below with the

basilar plate (text-fig. 35). A reconstruction of a rather earlier

stage seen from above (text-fig. 36) shows the capsule beginning

Text-fig. 37.

72° spg. mt.§

Left side view of a partial reconstruction of the anterior end. Cartilage is beginning

to appear. ‘The trabecule have been cut short. The pronephric funnels are

indicated in metaotic segments 5 and 6. The ventral portion of myotome 1

is still present.

Text fig. 38.

rade mt?

ET sos ee

Osa 4,

Fee,

seatetenmnneneentanwanees ct aeeee dacs g 0

“

Partial reconstructions seen from the right side. In text-fig. 88 ave the myotomes

and pronephric funnels. The trabeculae haye been eut short,

OF THE HEAD IN THE BATRACHIA URODELA. 109

to chondrify behind ; the occipital avch is fused on above the vagus.

The floor of the skull widens out considerably from the occipital

seoment forwards to allow space for the brain, and the preoccipital

arch is placed opposite the point where the occipital parachordal

plate meets the anterior parachordal plate, a point not yet chon-

drified. Although it appears to chondrify in continuity with the

auditory capsule, the preeoccipital arch is probably serially homo-

logous with the neural arches, as already suggested by Miss Platt.

Text-fig. 40.

Reconstruction, seen from above, of the head and anterior trunk-region. The dorsal

ends of the gill-arches are shown, also the ventral region of the myotomes.

Cranial nerve-roots are indicated on the right side.

Its first appearance as a procartilaginous rudiment is seen in text-

fig. 37, taken from a still younger larva, in the septum between the

2nd and lst myotomes. At this stage the ventral portion of the

first myotome (2nd somite) is still present. The auditory capsule

itself is scarcely recognisable and quite without cartilage.

In the next younger stage, text-figs. 38 and 39, the praoccipital

110 MR. E. S. GOODRICH ON THE OCCIPITAL REGION

arch has not appeared, the ventral portion of the first myotome is

larger, a ventral nerve-root is seen supplying myotome 2, and the

neural arches are mostly in a procartilaginous state.

Text-fig. 40 is a partial reconstruction of another larva of about

the same stage, but rather younger. The ventral portions of

myotomes | to 6 are shown, also the spinal ganglia. Procartilage

vaguely indicates the position of the auditory capsule, the occipital

arch is scarcely yet marked out, but the next three neural arches

Text-fig. 41.

= ite

hda

4 —

‘ \

er \ ‘ %

iy Cz. SO each

pin. gt.

gl. mt!

Reconstructions of the anterior region, seen from the right side. The visceral arches

and auditory sac are seen in text-fig. 41 ; these have been remoyed in text-

fig. 42, where the nerves are shown. ;

‘have begun to develop cartilage. The first myotome is quite,

and the second myotome nearly, cut into a dorsal and a ventral

portion by the roots of the glossopharyngeal and vagus. The

ventral remnant of the first myotome (2nd somite) varies con-

siderably in development, for in the younger larva drawn in

text-figs. 41 and 42 it is hardly distinguishable. At this stage the

trabeculee and visceral arches are the only skeletal elements visible,

and true cartilage can scarcely be said to have appeared.

OF THE HEAD IN THE BATRACHIA URODELA. 111

The next younger stage (text-fig. 43), again, shows a large

ventral piece of the first myotome. At a still younger stage

Text-fig. 43.

fF, as. mt fe?

Sg.

Left side view of a reconstruction of the anterior region.

The visceral arches are not completed.

(text-fig. 44), the trabecule have not appeared and the visceral

arches are represented by mere rods of procartilage.

Text-fig. 44.

mt.2 Gv. my /*+ as. fF,

Right side view of a reconstruction of the anterior end of a larva in which the

mesoblastic skeleton is represented only by procartilagimous rod-like visceral

arches,

We now come to embryos without true mesoblastic skeleton.

These earlier stages are most important in determining the number

of postauditory segments. Text-figs. 45 and 46 are reconstructed

from an embryo 5 mm. in length; they both are views from

above, but text-fig. 46 reaches farther down, so as to include

the alimentary canal and optic vesicles. Myotomes 4 and 5 are

WY MR. E. 8. GOODRICH ON THE OCCIPITAL REGION

clearly seen corresponding to the two pronephric funnels.

Myotomes 2 and 3 give off diverticula passing behind the 4th

branchial slit to form hypoglossal muscles. The ventral portion

of the first myotome is some way behind the auditory vesicle and

the rudiment of the hypoglossal ganglion. Below and in front

of the vesicle is the hyoidean somite; while in front of the

rudimentary spiracular gill-slit lies the mandibular somite,

Text-fig. 45.

Partial reconstruction, seen from above, of the anterior region of an embryo in which

the mesoblastic skeleton has not yet appeared. ‘The epidermal thickenines

corresponding to the glossopharyngeal and vagus nerves are imdicated by

dots.

Another embryo of about the same length, but a little younger,

is shown from the side in text-fig. 47. Here the auditory vesicle

is still quite continuous with the epidermis, and the relation of the

somites to the gill-slits is well shown. The slits are represented

OF THE HEAD IN THE BATRACHIA URODELA. eS

Text-fig, 46.

Similar reconstruction of the more dorsal region of the same embryo

as in text-fig. 45.

Text-fig. 47.

Left side view of a reconstruction of an embryo 5mm. long. The cavities of the

gill-pouches are shown in black. The auditory sac is a thickening directly

‘tS seontintious: with thelepiblast. Dots indicate epidermal.tilickenings, ©. © ii

Proc. Zoor, Soc.—1911, No. VITI. 8

114 MR. E. 8. GOODRICH ON THE OCCIPITAL REGION

in black, though not yet open. In front is seen the mandibular

somite, below the rudiment of the trigeminal ganglion. Behind

it lies the hyoidean somite, just in front of the vesicle and below

the facial rudiment. Posterior to the vesicle is the glossophar-

yngeal rudiment, below which extends the first metaotie somite

passing downwards into the first branchial arch. The vagus

rudiment extends over the next two and part of the 4th somite;

the first myotome is placed over the 2nd branchial arch.

Text-fig. 48 is a plan of a young embryo 3 mm. long, seen from |

the dorsal aspect, the nervous system being partly removed. The

Text-fig. 48.

Partial reconstruction of the anterior region of an embryo 3 mm. long, seen from

above. The greater part of the central nervous system has been removed to

expose the alimentary canal, notochord, &c. The first and second metaotic

somites overlie the first and second gill-arches.

position of the somite with relation to the other structures is

much the same as in text-fig. 47. Lastly, in text-fig. 49 is drawn

the youngest stage we need investigate. The dorsal portion of

the 2nd postauditory somite has become closely pressed on to the

3rd somite, with which it remains intimately associated. Indeed,

in the latter stages it is seen to be indistinguishably fused with

it, the two combining to form the anterior region of that dorsal

muscle which in combination with the 3rd myotome makes up

the temporal muscle of the adult. Neither.at this nor at any

OF THE HEAD IN THE BATRACHIA URODELA. 115

other stage does the first metaotic somite produce muscle-

substance.

Text-fig. 49.

Right side view of a reconstruction of an embryo 3 mm. long. The first metaotic

somite appears below the developing auditory sac and the glossopharyngeal

epidermal thickening. The 2nd somite is closely applied to the 3rd.

Conclusions.

From the foregoing account it will be understood that in

almost every particular my observations confirm the conclusion

reached by Miss Platt in her study of Mecturus. In Ambly-

stoma as in Necturus there are three occipital segments. The

first metaotic somite, however, disappears very early. Probably

the examination of an insufficiently complete series of stages

misled Sewertzoff (9) into the belief that there are only two

metaotic somites ; the first being either missed or later confused

with the second. Gaupp, in his excellent review of the develop-

ment of the skull (3), seems rather to favour Sewertzoff’s inter-

pretation. But if the diagram given by the latter, and repro-

duced by Gaupp, were correct, the second and third branchial

rami of the vagus would le outside the head area in the first two

trunk-segments ; and the third branchial ramus, passing behind

the last gill-slit, would then belong to the second trunk-segment

already provided with a complete spinal nerve possessing two

roots and a ganglion. This is obviously not the case. The

results of Sewertzoff and myself are compared in text-fig. 50.

Only in unimportant details do my own observations differ from

Miss Platt’s. For instance, I find a ventral nerve-root in seg-

ment 3 and a dorsal root in segment 5. Moreover, the ganglion

116 MR. E. 8. GOODRICH ON THE OCCIPITAL REGION

Text-fig. 50.

ee SONG

x &

AS)

ee ee ke Se a

Diagrams of the segmentation of the metaotic region in Amphibia :

B according to Sewertzoff, A according to my own observations.

of segment 4 seems to be early included in the rudiment of the

vagus ganglia. Finally, text-fig. 51 represents what I believe to

be the true composition of the Amphibian head.

Vext-fig. 51.

bbshe hypm

Diagram of the segmentation of the anterior region of an Amphibian.

We may now turn to the questions suggested on page 102. If

any segments have been suppressed, it 1s apparently behind the

vagus-root that they have been crushed out. Gegenbaur (4)

believed the occipital arch to bea compound of several- skeletal

OF THE HEAD IN 'THE BATRACHIA URODELA, 7

segments, not the homologue of a single neural arch as Stohr

supposed. Fiirbringer adopts much the same view in his great

monograph (2), concluding that the occipital condyle of the

Amphibia occupies the same segmental position as the last

occipital segment in the Selachii. Many segments are supposed

to have been crushed out between this and the vagus corre-

sponding to the region in the Selachian skull. through which

pass the spino-occipital nerves. He claims, indeed, to have found

possible traces of these segments in. Cryptobranchus, where he

discovered a small ventral nerve-root (z.) piercing the occipital

bone. Miss Platt and Gaupp (8 and 8) are inclined to accept

Fiirbringer’s view. Now the adoption by so many anatomists of

the theory that a number of segments have vanished from the

Amphibian head by a process of excalation is, I venture to think,

based not so much on facts as on theoretical considerations.

In the first place, there is a reluctance to admit that a structure

like the occipital condyle can be homologous in two groups,

although developed in different segments. But there can be no

doubt that the homology of an organ is independent of its

osition in the segmental series.

The hind limbs of a Frog, an Axolotl, and an Amphiuma are

homologous, in spite of the fact that they are placed in different

segments. It is unnecessary here to recapitulate in full argu-

ments which have already been given at length elsewhere (5, 6)

with regard to the development of the fins of fishes. But I may

briefly state :—That every trunk-segment is capable of producing

limb-elements; that is to say, of contributing to the formation of

median and paired fins. The shifting of a fin up or down the

body is not due to the migration of fin-material from one place to

another in the course of ontogeny; the fin, as a whole, arises

from that region of the trunk which it occupies in the adult (as

is shown by. the development and the nerve-supply). Relative

displacement in ontogeny is due almost entirely to “ concentra-

tion,” a relative narrowing of the base of the fin. Change of

position in phylogeny is brought about by progressive reduction

on one side, and growth on the other ; apparent migration is due

to certain segments beginning to contribute to the fin at one

end and certain other segments ceasing to contribute at the

other. By such “transposition” the fin may reach an entirely

new position.

Now, in the case of the fins of fishes, I have already shown (5)

that-it is not possible to account for variation in position by

the theory of inter- and excalation. Growth and transposition

from one segment to another alone account for the facts. The

same is probably true of the occipital condyle. There is not the

slightest trace of the disappearance of segments behind the vagus

in the ontogeny of the Amphibia. We are familiar with the

variation in the extent. of. the gill-region in.. Vertebrates. by mere

growth. Obviously the hind limit of “the series of gill-slitis varies

backward or forward, according: as certain segments. cease ta.

118 MR. E. 8S. GOODRICH ON THE OCCIPITAL REGION

develop gills or take on the function of gill-formation. The

posterior limit of the skull is doubtless altered in the same way,

and the position of the occipital condyles may shift up or down

the segmental series. There should, therefore, be no theoretical

objection to accepting the anatomical and embryological evidence

that the occipital region of the head in Amphibia contains only

three segments. If segments could really disappear, leaving no

trace behind, it would be hopeless to attempt to homologise

segments in any two forms.

There is another theoretical consideration which seems to have

led to the adoption of the view that the occipital region of the

Amphibia is not as simple as it appears. It is urged that if it

contained only three segments, the Amphibia would be more

primitive than the Fishes from which they have descended. The

possibility of the reduction in the number of occipital segments

has just been explained above; but is there really any necessity

to assume that it has taken place? From whatever Fishes the

Amphibia may have been derived, we may be sure it was not

from fully specialised Selachians. From paleontology alone we

may hope to obtain definite evidence on this point; until con-

trary evidence is brought forward, there is no necessity to assume

that the ancestors of the Amphibia had more than three differen-

tiated occipital segments. Of all the living fishes the Dipnoi are

those which most closely approach the Amphibia; even in the

modern Ceratodus there is no occipital condyle, no distinct limit

between head and trunk. It is true that several trunk-segments

have here been more or less completely assimilated to the skull

(Sewertzoff, 10); but there is no reason to believe that in the

remote common ancestor of the Dipnoi and Amphibia the dividing-

limit between the two regions could not have been developed

three segments behind the auditory capsule. This seems to be

in agreement with the conclusions of Agar (1), based on a study of

the development of Lepidosiren and Protopterus *.

The hypoglossus in the Amphibia and Amniota may certainly

be considered as homologous, although not necessarily composed

of the same segmental nerves. It is owing to the shortness of

the skull in the Amphibian that the hypoglossal roots come out

behind it.

Summary.

The chief contents of this paper may be summarised as follows.

Three occipital segments occur in the head of Amblystoma. Of

the three somites developed in these, the first forms no muscle

and disappears early ; the second forms a myotome divided into

dorsal and ventral portions, of which the former alone persists,

fusing with the myotome next behind. The myotome of the

third segment persists dorsally, that of the first trunk-segment

* The occipital region in fossil Amphibia seems to have been formed as in the

modern species. Important evidence as to the assimilation of the hypoglossal

segments in primitive Amniotes may perhaps be gleaned from a careful investiga-

tion of early fossil Reptiles. For instance, it seems to me not improbable that in

Paretasaurus the occipital region js still in an intermediate condition.

OF THE HEAD IN THE BATRACHIA URODELA. 119

combines with it to form the temporal longitudinal muscle

supplied in the adult by the first spinal nerve. The _ glosso-

pharyngeal nerve belongs to the first metaotic segment and has

no ventral root. To the second segment belongs the vagus root,

with which seems to be combined the ganglia of the next two

nerves. No ventral root was found in the second segment; but

a ventral root occurs in the last metaotic segment in early stages

of development. The first spinal issues from between the skull

and first neural arch as a ventral root only; it joins the second

spinal to form the hypoglossal nerve supplying muscles derived

from ventral outgrowths of the second, third, and fourth myotomes.

The basilar plate of the skull is formed by a backward growth of

the trabecule meeting a forward growth of the base of the

occipital arches. It fuses with the auditory capsules, and with

the preeoccipital arches developed in the septum between the first

and second myotomes. The occipital arches arise in the septum

between the second and third myotomes. There is no evidence of

the disappearance of segments behind the vagus, and no valid

objection to the view that the hind limit of the skull may shift

backwards or forwards in the course of phylogeny. At the same

time there is no reason to suppose that the ancestors of the

Amphibia had more than three occipital segments, when the

occipital condyle became clearly defined.

List of References.

1. Acar, W. E.—‘“* Development of the Anterior Mesoderm

and Paired Fins in Lepidosiren and Protopterus.” ‘Trans.

Roy. Soc. Edinburgh, vol. xlv. 1907.

. Firprincer, M.—‘ Ueber d. spino-occipitalen Nerven.”

Festschr. v. 1. Gegenbaur, vol. iii. Leipzig, 1897.

. Gauprp, H.—‘ Die Entwicklung des Kopfskelettes.” Hert-

wig’s Handb. Entw. Wirbeltiere, vol. ii. Jena, 1906.

. GEGENBAUR, C.—‘‘ Die Metamerie des Kopfes.” Morph.

Jahrb. vol. xiii. 1888.

5. Goopricu, KE. 8—‘‘ On the Development &c. of the Fins of

Fish.” Quart. Journ. Micr. Sci., v. 1906.

6. Goopricn, EH. 8.—The Vertebrata Craniata: Cyclostomes

and Fishes. Treatise on Zoology: part 9. London, 1909.

7. Parker, W. K.—‘‘On the Structure and Development of

the Skull in the Urodelous Amphibia.”—Part I. Phil.

Trans. vol. 167. 1876.

8. Puarr, J. B—“ The Development of the Cartilaginous Skull

and of the Branchial and Hypoglossal Musculature in

Necturus,” p. 377. Morph. Jahrb. vol. xxv. 1896-8.

9. Sewerrzorr, A.N. “ Die Entwicklung des Selachierschiidels.”

Festschr. C. v. Kupffer. Jena, 1899.

10. Sewerrzorr, A.N. “Zur Entwicklung des Ceratodus forsteri.”

Anat. Anz. vol. xxi. 1902.

11. Srour, Pa.— Zur Entwicklung des Urodelenschiidels,” p. 477.

Z. w. Gool. vol. xxxiii, 1880; “Zw: Entwicklung des

Anurenschiidels,” p. 68, /bid, vol. xxxvi. 1882.

Ee WwW ww

120 MR. OLDFIELD THOMAS ON THE MAMMALS.

Haplanation play ing on Text-figures..

ventral root.

spg., spinal ganglion.

a., auditory nerve. - ma., mandibular arch.

ac., auditory capsule. '-mdé., mandibular branch of trigeminal.

acp., ascending process of quadrate. mds., ., mesoblast of mandibular arch. -

as., auditory sac. mt., myotome 1-6.

bp., branchial plexus. mth., mouth.

br., branchial branch of vagus. mar., maxillary branch of abate Gl

brv., brain. | ma. 1, neural arch.

_brs., mesoblast of branchial arches —- zch., notochord.

1-4. ms., nasal sac.

brsl., branchial slit 1-4. 0e., occipital arch.

emm., median mandibular cutaneous | ocp., optic cup.

branch of facial. | opn., optic nerve.

col., columella auris. per., pericardium.

ctr., crista trabecule. | poe., preoccipital arch.

é.5 eye. | pop., preorbital process.

et., epiblastic thickening. | prf., profundus nerve.

Ff; facial nerve. | prn., pronephros.

Jac., floor of auditory capsule. | q-, quadrate.

foc., oculomotor foramen. | s.1,2,..., metaotic somite.

Sop. optic foramen. | sh., hyomandibular somite.

Gf» tacial ganglion and its epiblastic sk., hind limit of skull.

thickening. | sm., mandibular somite.

ggl.. ganglion of glossopharyngeal. | sop., superior ophthalmic nerve.

gl., glossopharyngeal nerve and its | 2 1, first spinal nerve.

epiblastic thickening. | sp.2d & ¥, second spinal nerve, dorsal and

gt., glottis.

gtr., ganglion of trigeminal and its

epiblastie thickening. | spl., spir acular slit.

gv., ganglion of vagus and its epi- spor., ventral root of spinal nerve.

blastic thickening. | tr., trabecula cranil.

h., hyomandibular branch of facial. | trg., trigeminal ganglion.

hda., hyoid arch. trp., trabecular parachordal.

hypm., hypoglossal muscle. ®., vagus nerve.

hyp., hypoglossal nerve. | tso., tectum synoticum.

im., internal mandibular branch of Silly, ventral lateral line nerve.

trigeminal. - vr., roots of vagus and glossopharyn-

j., anastomosis of Jacobson. | geal.

il., lateral line nerve. | vrt., ventral root.

m., medulla. {

9. The Mammals of the Tenth Hdition of Linnzeus ; an

Attempt to fix the Types of the Genera and the exact

Bases and Localities of the Species. By OLDFIELD

Tuomas, F.R.S., F.Z.S.

[Received November 29, 1910: Read December 13, 1910. ]

The tenth edition of Linné’s ‘Systema Nature’ is the recognized

basis and commencement of all systematic zoological work, but

doubtful questions in its interpretation are very ntimerous, and

there is probably nothing more desired, by mammalogists at least,

than a methodical examination of this important work, conah

suggestions, ontained on some definite and uniform system, for the

identification of its types, both of genera and species.

For many years I have taken a very great interest in this

subject, and have now ventured to prepare the present paper,

with the hope that it may be of service to other mammalogists Jess

favourably situated as regards books and specimens than I have

OF VHE TENTH EDITION OF LINN-EUS,. 421

the’ privilege of being. For in all the work I have. searcely in

a single instance been unable to obtain from the library of the

Natural History Museum what books I have wanted to verify

Linneus’s quotations, however rare and obscure they have been.

Without such an advantage it would have been impossible to

do the work at all.

With regard to the method used for finding the type-species of

the Linnean genera, I would draw attention to the very consider-

able extent to which the principle of tautonymy is available. In

a great proportion of cases Linnzeus selected as a generic name a

term which had been used by earlier writers for the single name

of a member of the genus, so that among the species of any given

genus there is commonly one which has ¢ among its synonyms the

name used by Linneus for the genus.

These names are generally quoted from Gesner, and comprise

the most conspicuous and best-known members of each genus, so

that in nearly all cases the genotype selected on this system is the

very one which by general consent we should wish to have for the

type. Any other rule that has been suggested, either “ first

species ” or “elimination,” is liable to produce results by no means

so satisfactory—indeed, the many objections to both are too well

known to need any insistence upon here. Nor can any later and

arbitrary selection of types without reason assigned be taken as

necessarily valid.

In two cases only, those of Simia and Dasypws, are results

arrived at different from those obtained by previous writers. The

shifting of the latter name from the rarer ‘‘ Huphractus” to the

really common Tatusia is unfortunate, but is necessary if we are

to get the benefit of the general stability which is obtainable from

the acceptance of the principle of tautonymy.

The case of Sima, however, is very different, and I confess to

hoping that the suggestions which are now being made for some

overruling decision in the case of certain well-known but

technically misapplied names may be carried out, both as toa

number of such names in general and to Sima in particular.

But it seems advisable, nevertheless, to put the technical case

irrespective of the prospect of any such decision, and therefore

(while hoping that a fiat attaching Simia to the Orang may go

forth) I have to point out that this name should be applied to

the Macaques, with S. sylvana (the Barbary Ape) as its type,

while Anthropopithecus will remain available a the Chimpanzees.

Should no such fiat as I speak of ever come to pass, this allo-

cation of Simia is at least better than its application to the

Chimpanzees. Not only is it less confusing in itself, owing to the

consequent total abolition of the combination Simia satyrus, con-

nected for a century with the Orang, but now sought to be applied

to the Chimpanzee, but also the genus to which it goes, that of the

Macaques, is already marked as the victim of a nomenclatural

change, for Pithecus antedates Macaca, and therefore one change

is not much worse than the other, pata hii eee se

122 MR. OLDFIELD THOMAS ON THE MAMMALS

Moreover, the Macaques are far less often dealt with or spoken

of by outside writers, not up to date in nomenclatural techni-

calities, than are the Anthropoid Apes, among whom such a change

has the maximum of inconvenience.

[After the completion of the present paper, there has come into

my hands a copy of the ‘‘ Opinions rendered by the International

Commission on Zoological Nomenclature,” dated July 1910, but

received by members of the Gratz Congress only at the end of

October.

In this publication the question of Linnean Genera in general

and Simia in particular is dealt with, certain conclusions are

come to not differing widely from my own, and the principle of

tautonymy as applied to the names in Linneus is on the whole

approved of.

Two genera (J/us and Camelus) have types definitely fixed for

them, twenty are mentioned as “seeming to retain as types” the

species I fix for them by tautonymy, though this is “not a ruling

by the Commission,” and Dasypus is ignored.

With regard to ‘Simia, the Commissioners, while seeing that

the rule brought the type on to sylvanus, would appear to have

thought it a benefit for zoology that satyrus should be considered

as such, an opinion in which I believe but few zoologists on

careful consideration would be found to agree with them. Under

this idea a brief but rather far-fetched and unconvincing argument

is adduced to avoid the simple conclusion about sylvanus, viz. :—

that “an examination of Gesner’s text shows that he did not

use Simia in the specific sense of ‘the simza,’ and that therefore

Simia Gesner, as used in the synonymy of S. sylvanws, cannot be

considered as the name of a species.”

Now I have carefully examined Gesner (three editions, including

that quoted), and can only say that his use of Simia is exactly

the same as his use of other names which are accepted as tau-

tonyms. On p. 847 he writes ‘“‘de Simia,” and gives a figure of

the Barbary Ape, and then on p. 855 he begins on other monkeys

under the heading “de Simiis diversis.” Exactly in the same

way we have “de Mure” on p. 714 (Mus musculus) and “ de

Muribus diversis” on p. 731; “de Mustela” on p. 752, “de

Mustelis diversis” on p. 762.

It is true that the Commissioners do not definitely accept as a

ruling of the Commission the above-mentioned twenty names on

the rigid basis of tautonymy, but seem to take as valid Palmer's

selection of the types as published in his Index (1904).

But the latter work cannot be accepted as that of a “ reviser”

in the true sense of the term. It is merely a bibliographical work

giving an indication of what species have been currently accepted

as types, without special revision by authors with all the facts

before them. In no case could I admit that it should override the

clear case for tautonymy presented by Stmia equally with the

twenty genera in which its results are agreed to both by Palmer

and the Commissioners |

OF THE TENTH EDITION OF LINNAUS. 123

The question of the species is not so simple as that of the

genera, having been complicated by the pages and pages which

commentators have thought it necessary to write balancing the

pros and cons in each case, weighing the importance ae this

reference or that, and deciding that such and such is the “ primary

quotation” and should be taken as the basis of the Linnean

name. This balancing method, however charming an exercise of

the writer’s ingenuity it may provide, is unfortunately full of the

personal equation, and therefore objectionable, while any method

which works automatically should be welcomed by those desirous

of coming to a final and impersonal decision on such points.

The system I advocate is that Linneus’s quotation of his own

earlier writings should be given an absolutely overriding im-

portance, and that, where these exist and can be traced back, all

others should be ignored.

This principle is not an arbitrary one, but is quite reasonable.

For when in 1758 Linneus gave a bmomial name to an animal to

which he had already given in 1748 a species-number and a

diagnosis (quoting it in the tenth edition), his idea of the species

would clearly have been formed at the earlier date, the later

edition only adding the binomial name. ‘To accept as of primary

importance in the determination of the latter some reference

which was not in existence at the time of the earlier edition

appears to me little short of ridiculous.

But if we take Linneus’s own Linnean quotations—which are,

in most cases, of the sixth edition—we get our field of selection

narrowed down at once to one or two easily traceable references,

from which the basis of the author's original idea of each species

is readily obtainable.

So important did Linnzus himself think the sixth edition that

practically every species of Mammal in it is quoted in the tenth,

and that merely by its page and number, as if that were the one

edition to which attention should be paid.

Curiously enough, this does not seem to be the case in other

groups than Mammals, the references to the sixth edition being

omitted, and primary stress laid on the ‘ Fauna Suecica.’

With regard to type localities, I have made every effort to

identify the places from which the specimens came that were the

original bases of Linnzus’s names. His own statements of

habitat are for the most part too vague to be of any use, while

accepting them literally would sometimes lead to grotesque errors,

such as the assertion, based on Linnzeus’s ‘‘ Hab. in Indiis,” that

“some Indian Armadillo” (!) should be taken as the basis of

Dasypus septemcinctus. “In Indiis,” in fact, hardly amounts to

more than a statement that the species is exotic.

But many localities can be settled from the authors quoted, as,

for example, in the case of Marcgrave, on whose descriptions *

Linneus’s names for a large number of the commoner Brazilian

* * Historie Rerum Naturalium,’ Brasilie, 1648.

124 MR. OLDFIELD THOMAS ON THE MAMMALS

species were based. With all these we are able to fix as a type

locality Pernambuco, for Marcgrave stayed at ‘“‘ Moritzstadt,’” now

Recife, most of his time, 1640-1644, while even his excursions

were limited to the coast region between 5° 45’ and 11°11’ 8. , thus

taking in little more than fr om Rio Grande do Norte to Alagoas,

a region at the centre of which Pernambuco lies.

Again, with regard to species named from the figures in Seba’s

‘Thesaurus,’ it would not be unjustifiable to suggest that i the

case of all tropical South American animals, Surinam—the great

source of all Dutch collections—should be accepted as the 1 type

locality, except when the preservation of the type specimen or

some incompatibility in the description shows this to have been

impossible.

The alterations in nomenclature that have proved to be necessary

are fortunately comparatively few, certainly far fewer than 1

had originally feared would be the case. Those not previously

published are the following :—

Simia. Genotype not satyrus but sylvanus. yp. 125.

Anthropopithecus satyrus for Chimpanzee. p. 125.

Technical name of Barbary Ape, Simia sylvana. yp. 125.

Pygathrix a distinct genus from Presbytis. p. 127 (footnote).

Macaca syrichta for M. philippinensis auct. _ p. 129.

Manis pentadactyla for M. dalmani Sand. p. 133.

Dasypus for Tatu; Huphractus tor “ Dasypus.” p. 141.

Arvicanthis striatus for A. pulchellus Gray. p. 148.

Physeter catodon for Ph. macrocephalus Linn. p. 157.

Other names here confirmed, which are unfamiliar, though

not now published for the first time, are those indicated for the

Whalebone Whales by Mr. F. W. True, those of certain monkeys

(Cebus, Mandzrill, &€.), recently pointed out by Dr. Elhot, Presbytis

aygula for P. mite atws, Cynocephalus volans for the Philippine

Colugo, Hlephas maximus for the Indian Elephant, and Luge meco-

phaya tridactyla for M. jubata.

' In one respect the present paper does not complete the exami:

nation of all the Linnean 1758 species, for a certain number aré

still unidentified pending the study of typical specimens formerly

in the ‘“*‘ Museum Regis Adolphi Friderici,” and no doubt in many

cases still in existence. I would commend to my colleagues in

Sweden an examination of the oid collections there preserved,

with an identification of such types as are still extant. Perhaps

Dr. Einar Lonnberg may be able to forward this final completion

of the study of the “ Zoological Bible ” written by bis immortal

compatriot.

In the following pages the genera and species are arranged

as in the ‘Systema,’ while opposite the name of each species on

the right-hand side of the page I have placed the name of the

recognized modern genus into which that species falls. U

OF THE TENTH EDITION OF LINNZUS. 125

Genus 1. Homo. p. 20,

Two species. Type, 7. sapiens.

Species :

1. sapiens. p. 20. Homo.

“« Nosce te ipsum.”

2. TROGLODYTES. p. 24.

A mixture of the abnormal hairy woman figured by Bontius

(Hist. Nat. Med. Ind. Or. p. 84) with aa account of albino negroes

taken from Nils Matson.

Genus 2. Simma._ p. 25.

Twenty-one species. Type by tautonymy: SS. sylvanus

(‘*Simia,” Gesn.).

The species are divided into three sections, (1) * Cauda nulla :

Simiz veterum, (2)** Cauda abbreviata: Papiones, and (3) ***

Cauda elongata: Cercopitheci.

From the name “Simi,” the type must clearly be sought

among the first section, which contains two species, satyrws and

sylvanus, and the second of these (not the jirst) should be taken

gi j the first, im fact—occurs

“Simia, Gesner, p. 847” (of 1620 edition, 957 of 1551 edition, p. 1

of German edition).

From this it would follow that, failing a fiat on the subject,

the generic name Stina should be transferred—not to the Chim-

panzee, as has been done by Matschie and Rothschild, but to the

Barbary Ape and its congeners.

Species :

1. ‘SATYRUS. Anthropopithecus.

Linnean reference: Syst. Nat. (6) p. 3, no. 1, where the first

quotation is

. Tulp. Obs. p. 270, pl. xiv.

Clearly the Common Chimpanzee*, whose name (again failing a

fiat) would therefore be Anthr opopithecus satyrus.

2. SYLVANUS. p. 25. (Sima or) Macaca t.

Linnean reference: Syst. Nat. (6) p. 3, no. 3, where the sole

reference 1s to

Alpin. Aegypt. p. 24], pl. xvi.

This is clearly a Barbary Ape, to which therefore, instead

* Of. W. Rothschild, P. Z.S. 1904, ii. p. 420.

+ I continue to use Micneuk soronia Saale pending a decision as to “ fiat” names.

To use Pithecus (although it antedates Macaca) would be merely the commence-

ment of a useless familiarity with that name, which would he agai ov ertlirow n

should the fiat*principle fail.

126 MR. OLDFIELD THOMAS ON THE MAMMALS

of Macacus (or Macaca) tnwus or Inwus ecaudatus, the name

Simia sylvana would appear to be technically applicable.

The reference Simia sylvanus has been already frequently put

among the synonyms of JMJacaca inua, so that the present identi-

fication is not a new one.

3. SPHINM, 20: (Mandril *.)

Linnean reference: Syst. Nat. (6) p. 3, no. 5, where the sole

reference is to Ray (Quadr. p. 158), who in turn speaks

of the “* Baboon Gesnero descripta et depicta.”

This is, no doubt, as shown by Dr. Elliot T, the Mandrill, of

which a figure and description are given on the page between 157

and 158 of Gesner’s German edition (1606).

4, APEDIA, p. 20. ‘)

Linnean (and sole) reference : Ameen. Acad.i. p. 278 (1749).

Not determinable from description, but if the specimen de-

scribed in the ‘ Ameenitates’ as being in the Museum Adolpho-

Fridericianum still exists, the species may yet be identified.

Type locality “in Indiis,’ which means hardly more than

“foreign.” No locality at all is mentioned in the ‘ Ameenitates.’

5. SILENUS. p. 26. Macaca.

No definitive reference, the only one, that to Alpinus (p. 242),

being with a query, so that the account should be treated by its

own evidence only.

The description and locality, however, appear to me to be

sufficient to assign the name to the Malabar “ Lion-tailed Monkey

of Pennant,” as was done by practically every author up to

Blanford.

That auther, working on the basis of the 12th edition, first

stated { that the name did not refer to the Lion-tailed Monkey,

and then later on used it for it §.

No exact typical locality available.

6. FAUNUS. p. 26. (2)

Linnean reference: Syst. Nat. (6) p. 3, no. 15, whence we

get Cercopithecus barbatus, no. 1, Clusius, Exot. p. 371

(1605).

Tt does not seem possible to determine this monkey with any

certainty. In the pointed white beard and white chest we have

something which suggests the Diana Monkey, while, on the other

hand, by the length of the beard (if mantle-hairs were mistaken

for a beard) and the tufted tail a Colobus of some sort might be

indicated.

No locality given.

* As Palmer has shown, this is the technical name for the Mandrill, if considered

generically distinct from other Baboons, as it apparently should be.

+ Ann. Mag. N.H. (8) iv. p. 417, 1909.

+ P.Z.S. 1887, p. 620. § Mamma. Ind. p. 16, 1888.

OF THE TENTH EDITION OF LINNAUS. lon

7. PANISCUS. p. 26. Ateles.

Linnean reference: Syst. Nat. (6) p. 3, no. 14, whence

Guariba, Marcer. Bras. p. 226.

Ateles paniscus.

Type locality: Pernambuco.

8. DIANA. p. 26. Cercopithecus.

Linnean reference: Act. Stockholm, 1754, p. 210, pl. vi.

Here a full description of the monkey is given, and, as Jentink *

has pointed out, it is the red-thighed form to which the name

diana was originally applied.

Type locality: “ Guinea.”’—Liberia (Jentink).

9. CEPHUS. p. 27. Cercopithecus.

No Linnean reference. Quotations of Muregrave (p. 228)

and Ray (p. 156).

Type locality : “ Guinea.”

10. ayeuLa. p. 27. Presbytis *.

No Linnean reference. Sole basis: ‘“‘Osb. iter. 99 ”(=Osbeck,

Ostindisk Resa, p. 99, 1757),

Type locality : Java.

The species usually known as P. mitratus Eschsch. (see Thos.

& Wrought. P.Z.8. 1909, p. 373).

11. HAMADRYAS. p. 27. Papio.

Linnean reference: Syst. Nat. (6) p. 3, no. 7, where the sole

reference is to Alp. Aegypt. p. 242, pls. xvii. & xviii.

Hamadryas Baboon.

Type locality : Egypt.

Te VACCHUSS en 2i(. Callithria..

Linnean reference: Syst. Nat. (6) p. 3, no. 12, whence Cagui_

minor, Maregr. Bras. p. 227 (misquoted 228).

Common Marmoset.

Type locality : Pernambuco.

13. @pipus. p. 28. Leontopithecus.

No Linnean reference. Sole basis : Edwards, Aves, iv. p. 195,

ple Ld. role

Type specimen said to have been brought from “ Vera Cruz, in

New Spain,” but the species does not occur there.

* Notes Leyd. Mus. xx. p. 233, 1898.

+ Dr. Elhot (Bull. Am. Mus. N.H. xxviii. p. 151, 1910) has recently pointed out

that the name Pygathria, based on Simia nemea, antedates Presbytis, and should

be used instead of it. But an examination of the skulls convinces me that nemea

and its close ally nigripes are, as Gray said, generically distinct from all the other

Langurs, which would therefore retain the name Presbytis. The basal axis of the

brain-case is set on to the facial bones at quite a different angle in ‘the two genera,

128 MR. OLDFIELD THOMAS ON THE MAMMALS

14, mreiors. p. 28. i Cercopithecus.

Linnean reference: Linn. in Hasselquist, Iter, p. 190, 1757.

Formerly known as C. griscoviridis.

See de Winton, in Anderson, Mamm. Egypt, p. 15, 1902.

Type locality : Upper Egypt.

15. Mipas. p. 28. Leontopithecus.

No Linnean quotation. First and primary reference :

Edwards, Aves, iv. p. 196, pl. 196, 1751.

Locality : ‘“‘ West Indies.”

16. cyNAMOLGOS*. p. 28. Papio.

Linnean references: Syst. Nat. (6) p. 3, no. 10 (ex Maregr.

Bras. p. 227), and Simia Pay place Linn, in Hasselq. Iter,

p- 189, 1757.

Both these references being Linnean, the one that esters to an

actual specimen should be faken in preference to that merely

based on Maregrave’s short and indefinite notice.

Consequently Sima cynamolgos becomes. a synonym of Papio

hamadryas.

Type locality : Upper Egypt.

17. APELLA. p. 28. Cebus.

Linnean reference: Mus. Ad. Frid. p. 1, pl. 1., 1754.

Cebus capucinus auctorum.

See Elliot, Bull. Am. Mus. N. H. xxvi. p. 227, 1909.

No type locality available, but Guiana may be accepted from

later authors.

18. morta. p. 29. Saimiri.

Sole basis: Seba, Thes. 1. p. 52, pl. xxxiii. fig. 1, 1734.

Clearly a Squirrel-Monkey, but as the species is not determinable

the name must lapse. ,

The suggestion that Surinam should be taken as a typical

locality for Seba’s animals cannot be adopted in this case, as the

statement that Samia morta had the back of the head blackish is

inconsistent with the condition found in the Guianan species.

19: cAPUCINA. p. 29. Cebus.

Linnean reference: Mus. Ad. Frid. p. 2, pl. ii., 1754.

The monkey usually known as C’. hypolewcus Humb.

See Elliot, Bull. Am. Mus. N. H. xxvi. p. 227, 1909.

No type locality available. The species is Central American

and Colombian.

being far more strongly inclined in Pygathrix, in which, as a consequence, the

posterior nares are of enormously greater height. Gray, following Dahlbohm,

‘separated the two on external characters and the proportions of the fore and hind

limbs, and my comparison of the skulls quite confirms his judgment i in the matter.

“*® Spelling altered to cynomolgus in 12th edition. 9 he en

OF THE TENTH EDITION OF LINN EUS. 129

20. SCIUREA. p. 29. Saimiri.

Linnean reference: Mus. Ad, Frid. i. p. 3, 1754,

Common Squirrel-Monkey.

The Guianan form, which agrees with the description, may be

accepted as typical.

21. syRICHTA. p. 29. Macaca.

Linnean reference: Syst. Nat. (6) p. 3, no. 6; whence we get

“‘ Cercopithecus Luzon. minimus D. Camel,” Petiver, Gazyo-

phylacium, pl. xiii. fig. 11. No.176 of Index.

This figure—although a ghastly travesty in itself—can be

nothing but the one monkey found in Luzon, viz., Macaca

philippensis, philippinensis, or palpebrosus auctorum, which names

should give way to that of WZ. syrichta.

Type locality: Luzon (probably Southern Luzon, whence

Dr. Camel sent “‘ Lemur volans”).

Genus 3. Lemur. p. 29.

Three species, no one with “ Lemur” as a subsidiary name.

Type, Z. catta; L. tardigradus having been made the type of

Loris, Geoff., 1796, and £. volans of Cynocephalus, Bodd., 1768

(= Galeopithecus, Pall., 1780).

All authors have aecepted ZL. catta as the type of Lemur.

Species :

1. TARDIGRADUS. p. 29. Loris.

Linnean references: “‘ Mus, Ad, Fr, i. p, 3; Syst. Nat. (6)

Pao NO. 2.

The specimen referred to in Mus. Ad. Frid. and redescribed in

the 10th edition is still in the Stockholm Museum, and, as

Dr. Lénnberg informs me, is the Cinghalese Slender Loris. It

may even have been the specimen figured by Seba, who is quoted

in the 6th edition.

See also Thos. Ann. Mag. N. H. (8) i. p. 467, 1908, where the

same conclusion 1s reached on the ordinary ‘‘ Linnean reference ”

grounds.

Type locality : Ceylon.

2. cATrA. p. 30. Lemur.

Sole basis: Simia—Sciurus madagascariensis, sive Maucauco,

Edwards, Aves, 119, t. 199.

The Ring-tailed Lemur.

3. VOLANS. p. 30, Cynocephalus.

References :—

Cato—Simius volans camelli, Pet.(iver), Gaz.(ophylacii

Nature et Artis), t. 9. f. 8 (1702); Aet. Angl. 277,

* Misprint for 3.

Proc. Zoou. Soc.—1911, No. IX. 9

130 MR. OLDFIELD THOMAS ON THE MAMMALS

p. 1065 (= Phil. Trans. Roy. Soe. pt. 277, included in

vol. xxii. 1704).

Vespertilio admirabilis, Bont.(ius), Java, p. 68, 1658.

Felis volans ternatia, Seba, Mus. i. p. 93, t. 58. f. 2, 3.

The first is the primary and only vital reference, as the others

refer to specimens from impossible localities (Guzerat and Ternate).

Petiver’s type locality was Pampanga, Southern Luzon, and the

animal therefore was that until recently known as G'aleopithecus

(or Colugo) philippinensis.

Cynocephalus, Boddaert, was based on “ Lemur volans Linn.,”

and antedates Galeopithecus by many years.

See Thos, Ann. Mag. N. H. (8) i. p. 252, 1908.

Genus 4. VESPERTILIO. p. 31.

Seven species, of which no. 7, marinus, has among its synonyms

Vespertilio, Bell. Av. 147; Gesn. Av. 694.

Therefore type by tautonymy: Vespertilio murinus L., the bat

formerly known as V. discolor.

See Miller, Ann. Mag. N. H. (6) xx. p. 379, 1897.

Species :

1. vAMPYRUS. p. 31. Pteropus.

Linnean reference: Syst. Nat. (6) p. 7, no. 1.

There the only reference is to

Canis volans ternatanus orientalis, Seba, Thes. i. p. 91; t. 57.

We diy Oe

Pteropus edulis auctorum.

See Thos. P. Z. 8. 1892, p. 316.

Type locality: Ternate. A co-type in British Museum.

2, SPECTRUM. p. 3l. Vampyrus.

Only reference: Canis volans maxima aurita, Seba, Thes, i.

10s SAG te Bley ey Il

Species known rightly as Vampyrus spectrum L.

Suggested type locality : Surinam.

3. PERSPICILLATUS. p. 31. Hemiderma.

Linnean references: Syst. Nat. (6) p. 7; Mus. Ad. Fr. i.

Ds Us

The latter, as referring to an actual specimen, would’ take

precedence, but the description is unrecognizable, and no specimen

is now in existence, as Dr. Lonnberg informs me. We therefore

take the single reference of the 6th edition :—Vespertilio

americanus vulgaris, Seba, Thes. 1. p. 90, t. 55. f. 2, of which the

original specimen (B.M. no, 67.4.12.597) is the bat known until

1892 as Hemiderma brevicauda.

See Thos. P. Z. 8. 1892, p. 316.

Suggested type locality: Surinam.

OF THE TENTH EDITION OF LINNUS. Si

4, SPASMA. p. 32. Megaderma.

The Linnean reference (Syst. Nat. (6) p. 7) refers alone to

Glis volans ternatanus, Seba, Thes. i. p. 90, p. 56. f. 1, which

is the bat known as

Megaderma spasma.

See K. Anders. & Wrought. Ann, Mag, N, H. (7) xix. p. 132,

1907.

5. LEPORINUS. p. 32. Noctilio.

Equally through Syst. Nat. (6) p. 7, the first Linnean refer-

ence, and Seba, pl. lv. fig. 1, we get to the correctly known

Noctilio leporinus.

Suggested type locality : Surinam.

6. AURITUS. p. 32. Plecotus.

References: Frisch, Av. (Vogel Deutschlands) 103.

Kdw. Av. t. 201. f. 3.

Olear. Mus. t. 15. f. 1.

The Long-eared Bat, Plecotus auritus,

Type locality, from Frisch : Germany (Berlin).

7. MURINUS. p. 32. Vespertilio.

Linnean references: Faun, Suec. 18; Syst. Nat. (6) p. 7, no. 2.

For reasons explaining why this should be considered as the bat

formerly known as Vespertilio or Vesperugo discolor, see Miller, 1. c.

Type locality: Sweden (Upsala).

Genus 5. ELEepHas. p. 33,

Monotypic. Type, #. maximus (H. indicus auctorum).

Species :

MAXIMUS. p. 33. Llephas.

Linnean reference: Syst. Nat. (6) p. 11, where the single quo-

tation is to Ray, Quadr. p. 123 (131 in the Museum copy).

Ray gives a long account of the Elephant, but does not

distinguish the two forms, nor give any locality. Linneus,

however, says ‘ Habitat in Ceylone paludosis,” whence

Type locality : Ceylon.

Genus 6. TRicnEcuUs. p. 34.

Monotypic. Type, 7’. manatus.

Species :

MANATUS. p. 34. Trichechus.

Linnean reference: Syst. Nat. (6) p. 39 (misprinted 37).

Single reference in 6th edition: Artedi, Synonym. 109

(misprinted 107).

Artedi here, again, gives a primary reference to his own ‘ Genera

Kaw MR. OLDFIELD THOMAS ON THE MAMMALS

Piscium,’ no. li. p. 556, where we find a general account which

includes both Manatees, American and African, and Dugongs.

Linnzus, however, puts ‘in mare americano” as locality, and this

would fix the name on the common American species.

Type locality : West Indies.

Genus 7. Brapyeus. p. 34.

Two species. Type by selection: B. tridactylus. Selected by

Illiger, who at the same time eliminated B. didactylus as a

Cholepus.

Species :

1. TRIpDAcTYLUS. p. 34. Bradypus.

First Linnean reference : Syst. Nat. (6) p. 3.

Two references : Seba, Thes. i. t. 33. f. 2, and Maregr. Bras.

221, of which the first may be taken as primary.

Type locality (suggested): Surinam.

2. DIDACTYLUS. p. 35. Cholepus.

First Linnean reference: Syst. Nat. (6) p. 3.

One reference only : Seba, Thes. i. t. 34.

Type locality (suggested): Surinam.

Genus 8. MyrmrcopHaea. pp. 35.

Three species. Type by elimination : J/. tridactyla, M. didactyla

having been taken out as Cyclopes and WM. tetradactyia as

Tamandua.

Species :

1, DipAcTyLA. p. 35. Cyclopes.

Based on specimen. Fuller description Mus. Ad. Frid. p. 8.

No locality, but as an identification is made, through Syst.

Nat. (6) p. 8, with Seba, pl. 37. fig. 3, we may take as the

Type locality : Surinam.

2. TRIDACTYLA. p. 35. Myrmecophaga.

Linnean reference: Syst. Nat. (6) p. 8, where we find “

Quadr. 241. Marcer. bras. 225.”

Ray’s account simply refers to, and is based on, that given by

Marecgrave of the Tamandua guacu, the Great Anteater, of which

he gives a recognizable figure.

The Great Anteater, commonly known as Myrmecophaga jubata,

should therefore, as pointed out in 1901, be known as I. tri-

dactyla.

Type locality: Pernambuco.

See Thos. American Naturalist, xxxv. p, 148, 1901,

Ray,

OF THE TENTH EDITION OF LINN EZUS. 133

3. TETRADACTYLA. p. 35. Tamandue,

Linnean reference: Syst. Nat. (6) p. 8, which gives “ Ray,

quadr. 242. Marcgr. bras. 226.”

Here, again, as in the last species, Ray simply refers back to

Marcgrave, whose animal is the Tamandua.

Hence the common name Tamandua tetradactyla is correct.

Type locality: Pernambuco.

Genus 9. Manis. p. 36.

Monotypic. Type, I. pentadactyla.

Species :

PENTADACTYLA. p. 36. ; Manis.

Linnean reference: Syst. Nat. (6) p. 8, where four quotations

are given :—

Bontius, Ind. 60 (1658).

Clus. exot. 347.

Olear. Mus. 8, t. 7. f. 1.

Ray, Quadr. 247.

Of these, Bontius, as being the first quoted, should be taken ;

and here we find the actual locality mentioned, namely, Tajoan =

Tai-wan = Formosa.

On this account the species which has ordinarily borne the

name of JM. dalmani Sund., should be considered as the true

M. pentadactyla of Linneus.

Moreover, Linnzus also includes in his synonymy Dalman’s

paper (Act. Stockh. 1749, p. 265) on a Pangolin from China or

Formosa ; and as he must have seen the specimen when he gave

the name pentadactyla, the propriety of the above identification

is confirmed.

Type locality : Formosa.

Genus 10. PHoca. p. 37.

Four species. Type by tautonymy: PA, vitulina (“‘ Phoca seu

Vitulus marinus,” Gesner).

Species :

1. unsinA. p. 37. Callorhinus.

Sole reference : Steller, Nov. Comm. Petrop. ii. p, 331 (1751).

The Fur-Seal, commonly known as Callorhinus ursinus.

Type locality : Bering Island.

2. LEONINA, p. 37. Mirounga.

Sole reference: Anson, Itin. p. 100.

Sea-Lion.

Type locality: Juan Fernandez.

134 MR. OLDFIELD THOMAS ON THE MAMMALS

3. ROSMARUS. p. 38. Odobenus.

Linnean quotation : Syst. Nat. (6) p. 6, no. 2 (inisprinted 8).

Correctly known as Odobenus rosmarus (not Odobeenus, if

Brisson is taken as the founder of the generic name).

Type locality: Northern Atlantic.

4, VITULINA. p. 38. Phoca.

Linnean reference: Syst. Nat. (6) p. 6, no. 1, which refers

again to Fauna Suecica, no. 11 (p. 4).

Common Seal.

Type locality : “in Mari Bothnico et Baltico.”

Genus 11. Canis. p. 38.

Seven species. Type by tautonymy: Canis familiaris, among

whose synonyms there appears “ Canis Gesner.”

Species :

1. FAMILIARIS. p. 38. Canvis.

Linnean references: Faun. Suec. no. 125 Syst. Nat. (6) p. 5.

Type locality : Sweden (Upsala).

2. LUPUS. p. 39. Canis.

Linnean references: Faun. Suec. no. 13; Syst. Nat. (6) p. 5.

Type locality : Sweden.

3. HymNA. p. 40. Hyena.

Linnean reference: Syst. Nat. (6) p. 5, where the single

reference is to Kempf. Ameen. p. 407, fig. 4 (letterpress

on p. 411).

The animal commonly called Hyena striata, but whose proper

name is Hyena hyena.

Type locality: Bennt Mts., Laristan, S$. Persia, Kempfer

also mentions a specimen from Isfahan,

4. vuLpEs. p. 40. Vulpes.

Linnean reference: Syst. Nat. (6) p. 5, no. 6.

Correct name: Vulpes vulpes.

Type locality : Sweden (Upsala),

5. ALOPEX, p. 40. (Vulpes.)

Linnean reference: Syst. Nat. (6) p. 5, no. 5, where the

whole entry is

“Canis cauda recta extremitate nigra G'mel. Vulpes cam-

pestris.”

I cannot identify this animal. It has been supposed to be an

individual variety of V. vulpes, which Blanford (P.Z.S. 1887,

p. 635) proposed should bear the name J’. alopen.

OF THE TENTH EDITION OF LINNAUS. 135

6. LAGOoPUS. p. 40. Alopex.

Linnean reference: Syst. Nat. (6) p. 5, where V. alba and

V. cerulescens are quoted from Kalm and the ‘ Fauna

Suecica.’

Arctic Fox.

Type locality: Sweden (Lapland).

7. AuREUS. p. 40. Canas.

No Linnean reference, that to the 6th edition being omitted.

However, the first reference, Kempf. Ameen. p. 413, fig. 3,

is the one quoted in that edition, p. 5, no. 5.

Jackal.

Type locality: Bennda Mts., Laristan, S. Persia.

Genus 12. Feuis. p. 41.

Seven species. Type by tautonymy:; Felis catus (‘‘ Felis” Aldr.),

Species :

1. neo. p. 41. ‘elis.

Linnean reference: Syst. Nat. (6) p. 4, no. 1, where the

quotation runs “ Leo. Dod. gall. t.1& 17.” The plate 1

referred to appears to be that in Mém. Acad. Sci. Paris, iil.

1666-1699, p. 1, pl.1., publ. 1733, where there is an account

of the dissection of four lions and three lionesses.

No type locality available.

2. TIGRIS. p. 41. Felis.

Linnean reference: Syst. Nat. (6) p. 4, no. 2, where two

quotations are given :—

Ray, Quadr. p. 165; Bont. Jav. p. 53.

Taking the first of these, we find a description which mixes up

the Jaguar with the Tiger, but Linné’s “ Habitat in Asia ” may

be taken as restricting the name to the Tiger.

Later revisers of the races of Tigers, of whom Fitzinger and

Matschie may be specially quoted, have restricted F. tigris to

India proper, and particularly Bengal.

3. PARDUS. p. 41. Felis.

Linnean reference: Syst. Nat. (6) p. 4, no. 3, where we find

Ray, Quadr. p. 166, and Alp. Aegypt. p. 237.

Ray gives no indication of locality, but passing to Alpinus we

find an account of Leopards seen at Cairo and Alexandria alive in

captivity. The account is, no doubt, partly based on Hunting

Leopards (Cynclurus), but, none the less, may be accepted as

giving for the

Type locality: Egypt.

136 MR. OLDFIELD THOMAS ON THE MAMMALS

It may be here reiterated that Linne’s “in Indiis” cannot be

taken to mean India in the modern sense, but means hardly more

than ‘‘from abroad.” In this case, he gives a synonymy based on

African and Mexican references, and then simply puts ‘* Habitat in

Indiis.”’

4, oncaéA. p. 42. Felis.

First reference: Ray, Quadr. p. 168; and this in turn is

based primarily on Marcgrave’s account of the Jaguar.

Consequently

Type locality : Pernambuco.

5. PARDALIS. p. 42. Felis.

Linnean reference: Syst. Nat. (6) p. 4, no. 4, where the only

citation is

“ Catus pardus, Hernandez, Mex. p. 512.”

Ocelot.

Type locality: Mexico.

Linneus also saw and described as the same species a living

specimen “in Vivario cliffortiano,” but there is nothing in this

description inconsistent with the taking of the Mexican Ocelot as

the typical form.

6. catus. p. 42. Felis.

Linnean references: Faun. Suecica, no. 3; Syst. Nat. (6)

p- 4, no. 6.

Clearly the ‘‘ Blotched Tabby ” Domestic Cat.

See Pocock, P. Z.S. 1907, p. 149.

Type locality : Upsala.

7. LYNX. p. 43. Lynn.

Linnean references: Faun. Suecica, no. 4; Syst. Nat. (6)

p. 4, no. 7; It. Wgoth. p. 222.

North European Lynx.

Type locality: Wennersborg, 8. Sweden. (From the first

mention in the Westgotha Resa.)

Genus 13, VIvERRA. p. 43.

Five species.

Type, both by elimination and selection: V. zivetha.

Species:

1. ICHNEUMON. p. 43. Mungos.

Linnean references: Linn. in Hasselq. Itin. p. 191, 1757, and

Syst. Nat. (6) p. 6, no. 3.

Common Mongoose.

Type locality : Egypt.

OF THE TENTH EDITION OF LINN.ZUS. 137

2. MmMPHITIS. p. 44. (2)

No Linnean reference.

“Ysquiepatl, Hern. Mex. 332; Seb. Mus. i. p. 68, pl. xhi.

fig. 1; Rai, quadr. p. 181.”

It does not seem possible to determine to what animal this

name should be applied. Hernandez gives a figure which has a

superficial resemblance to a Tayra, but the description of the

tail is quite inconsistent with this idea, and is applicable rather

to aSkunk or Bassariscus. Seba’s animal was probably a Vasua.

Linneus himself says in one place that the colour is brown

above and yellowish below, and in another that it is white above

and variegated black and white below, an inconsistency which

justifies one in treating the name as unrecognizable.

3. PuToRIUS. p. 44. Spilogale.

No Linnean reference.

“ Putorius americanus striatus, Catesb. Carol. ii. p. 62.”

Alleghenian Spotted Skunk.

Type locality : South Carolina.

See Howell, N. Am. Faun. no. 26, pp. 15-17, 1906.

4, ZIBETHA, p. 44. Viverra.

No Linnean references, that in the 6th edition, p. 6 (Meles,

no. 2), being omitted in the 10th. It quotes “ Ray, Quadr,

178; Dod. gall. 155.”

General references to Clusius, Ray, Dodart, Hernandez,

Olearius, and Beltonius.

From the name zibetha we may take the “ Animal zibethicum’

of Ray, Dodart, Hernandez, and Olearius as the primary reference,

and of these we should take the first in Ray, who is also the first

quoted in the omitted 6th edition reference.

Turning to Ray we find he quotes the description by Faber in

Hernandez, Nov. Hispan. pp. 538, 580, and 581 (also quoted by

Linneus). Taking, again, the first of these we find an animal de-

seribed which has a ringed tail, and would therefore be an Indian

Civet. The illustration does not show the rings properly, but

they are sufficiently indicated in the description. All the other

references are to the African V. civetta.

Type locality (from Faber): Bengal.

5, GENEITA. p. 45. Genetta.

No Linnean reference.

“Genetta s. Ginetta, Rai, quadr, 201. Bell. itin. 76. Gesn,

quadr. 550.”

The first reference in Ray being to Spain, that may be taken as

the type locality, especially as the basis of the name (Gineta)

remains as the Spanish colloquial term for the animal,

138 MR. OLDFIELD THOMAS ON THE MAMMALS

Genus 14. Musrena. pp. 45.

Nine species.

Type by tautonymy: J. erminea (‘* Mustela” Gesner ; mis-

quoted Mustela vulgaris).

Although in this instance alone of the Gesner quotations of the

‘Systema’ a second name is attached to the primary one, for which

tautonymy is claimed, this proves to be a misquotation by Linnzus,

and the correct quotations seem actually to strengthen the case

for the Ermine being really the typical tautonymous species. For

though Linneeus quotes ‘ Mustela vulgaris, Gesner,” we find, as a

matter of fact, that Gesner* did not put Mustela vulgaris, but

simply ‘“ Mustela” in the German edition (that quoted in the

1758 ‘Systema ’) and in the Latin one *‘ Mustela proprie sie dicta,”

in contradistinction to his “ A/ustelis diversis,” which included the

Marten and Polecat.

The Ermine was therefore clearly the Mustela of both Gesner

and Linneus, and should be treated as the type species.

Species :

1, Lurris. p. 45. Latax,

No Linnean reference.

Lutra marina, Act. Petrop. 1749, p. 267, 7. e. Steller, Nov.

Comm. Petrop. ii. p. 367 (1751).

Lutra brasiliensis, Rai, quadr. p. “ 187” (189).

The primary reference is clearly to Steller, and the type

locality Kamtchatka.

2. LUTRA. p. 45. Lutra.

Linnean reference: Faun. Suecica, no. 10.

European Otter.

Type locality : Upsala.

3. GULO. p. 45. Gulo.

Linnean reference: Faun. Suecica, no. 6.

Glutton.

“ Habitat in alpibus Lapponicis, Dalekarlicis, sylvisque

vastissimis.”

Taking the first place mentioned we have

Type locality : Lapland.

4, BARBARA. p. 46. Tayra.

Tayra, by description.

“ Habitat in Brasilia, M. Ac. Holmens.”

* The pagination of Gesner quoted by Linneus is somewhat confusing. While, in

the majority of cases (e. g., the Putovius), the pages are quoted from the 1620 Latin

edition, the “ Martes” and the “ Mustela vulgaris” are given as “151” and “ 149,”

these pages coming from the German edition of 1606. Their respective numbers in

the 2nd Latin edition (1620) are 764 and 752, and in the original edition (1551) 865

and 801.

OF THE TENTH EDITION OF LINNAEUS. 139

5, MARTES. p. 46. Martes,

Linnean references: Faun. Suecica, no. “3” (7); Syst. Nat.

(6) p. 5, no, 2.

Pine Marten.

Type locality : Upsala.

6. PUTORIUS. p. 46. Mustela.

Linnean references: Faun. Suecica, no. 8; Syst. Nat. (6)

p. 9, no. 3.

Polecat.

Type locality: Scania, S. Sweden.

7. FURO. p. 46. Mustela.

No Linnean reference.

“* Mustela sylvestris, Viverra dicta, Rai, quadr. 199.”

Ferret.

‘Said to be from Africa.”

8. ZIBELLINA. p. 46. Martes.

No Linnean reference.

“* Mustela s(z)ibellina, Rai, quadr. p. 201.”

Sable.

Type locality: N. Asia,

9. ERMINEA. p. 46. Mustela.

No Linnean reference.

“‘ Mustela vulgaris, Gesn. quadr. 149 ” (German edition).

Stoat or Ermine.

Type locality ; “‘ In unseren Landen” (Germany).

Genus 15. Ursus. p. 47.

Four species. Type by tautonymy: U. arctos (‘“ Ursus,

Gesner”).

Species :

1. arctos. p. 47. Ursus.

Linnean references: Faun. Suecica, no. 2; Syst. Nat. (6)

p- 4, no. 1,

European Brown Bear,

Type locality (ex Faun. Suec.): Northern Sweden.

2. Luscus. p. 47. Gulo.

No Linnean reference.

Ursulus lupo affinis americanus, Edw. av. ii. p, 1038, t. 103,

1747.

American Glutton.

Type locality : Hudson’s Bay.

140 MR. OLDFIELD THOMAS ON THE MAMMALS

3. MELES. p. 48. Meles.

Linnean references: Faun. Suec. no. 15; Syst. Nat. (6) p. 6.

European Badger.

Type locality : Upsala.

4. Loror. p. 48. Procyon.

Linnean references: Syst. Nat. (6) p. 4; Act. Stockh. 1747,

bei Loa (DERI aseqae).

Although in the last-named place Linneus refers to earlier

accounts, yet his description of an actual specimen, whose habit,

as mentioned by him (p. 282), of dipping its food in water would

have suggested the name lotor, should override his synonymy.

His statement that the Raccoon is called Ispan by the Swedes

in Pennsylvania may be accepted as defining the locality of his

animal. Therefore

Type locality : Pennsylvania.

Genus 16. Sus. p. 49.

Four species. Type by tautonymy : Sus scrofa (Sus, Gesn.”).

Species :

1. sorora. p. 49. Sus.

Linnean references: Faun. Suec. no. 36; Syst. Nat. (6) p. 12.

Kuropean Wild Swine.

Type locality : Germany, whence the wild specimens referred to

by Linneus were introduced * into the island of Oeland.

2. PORCUS. p. 90. Potamocherus.

Linnean reference: Syst. Nat. (6) p. 12.

Refers to “‘ Rai, quadr. 96,” where a further reference is given

to Maregrave’s Porcus guineensis, whose description is clearly

that of the West-African River-Hog.

Type locality : Guinea,

SU ATACU. a pa. Tayassu.

Linnean reference: Syst. Nat. (6) p. 12, no. 3.

Here the only quotation is Ray (Quadr. p. 97), who in turn

gives primary importance to Tyson’s description (Phil. Trans. Roy.

Soc. xii. p. 359, 1683) of what he calls a Mexican Musk-Hog.

We may therefore consider the

Type locality: Mexico.

4, BABYRUSSA. p. 90. Babirussa.

Linnean reference: Syst. Nat. (6) p. 12, no. 4, where we find

two quotations : Ray, Quadr, p. 96, and Seba, Thes. 1. p. 80,

pl. 1. figs. 2 & 3.

Babirusa.

Type locality (from both Ray and Seba): ‘‘ Island of Boero.”

* See Lilljeborg, Sver. Rygg. Dagedj, pt. ii. p. 761, 1874.

OF THE 'TENTH EDITION OF LINN-EUS. 14]

Genus 17. Dasypus. p. 50.

Six species. Type by tautonymy: D. novemeinctus (“ Dasypus,

Hern.”).

Apart from the difficult and doubtful case of Sima, this is the

only instance in which the use of tautonymy in selecting Linnean

types brings out a result contrary to common use.

Linneus placed the name ‘‘ Dasypus” among the synonyms of

D. novemcinctus alone, and the account 1 Hernandez from which

this name comes is clearly that of one of the group long known as

Tatusia, more recently as Tatu.

The comparatively recent date of the latter change, the extreme

ugliness of Z'atu, which antedates Tatusia, and the classical suita-

bility of Huphractus, which stands for seacinctus and its allies,

will all help to reconcile us to the shifting of Dasypus from one

group to another.

This shifting is quite unavoidable, if the invaluable principle of

tautonymy is to be utilized at all.

Species :

1. unicrnctus, p. 50. Cabassous.

Linnean reference : Syst. Nat. (6) p. 6, no. 4, where the sole

reference is to Seba, Mus. 1. p. 47, pl. xxx. figs. 3 & 4.

This, or at least fig. 3 (fig. 4 being more doubtful), is clearly the

Tatouay or Twelve-banded Armadillo.

No original locality available, but being from Seba, Surinam

may be suggested, if Cabassows occurs there, which is not as yet

certainly known. t

2. TRICINCTUS. p. 51. | Tolypeutes.

Linnean reference: Syst. Nat. (6) p. 6, no. 2, where the sole

quotation is to Seba, 1. p. 62, pl. xxxvilil. figs. 2-3.

Seba’s animal is a typical Three-banded Armadillo.

Suggested type locality : Surinam.

3. QUADRICINCTUS. p. 51. Tolypeutes.

Linnean reference: Syst. Nat. (6) p. 6, no. 3, where the single

quotation is to “Column. aquat. 11. p. 15. pl. xvi.” (1606).

Colonna’s animal is a four-banded specimen of Tolypeutes

tricinctus, of which D. quadricinctus is therefore a synonym.

4. sSEXCINCTUS. p. 51. Euphractus.

Linnean references: Mus. Ad, Frid. p. 7 (1754), and Syst.

Nat. (6) p. 6, no. 5.

The former takes precedence, as being based on a specimen,

whatever the reference to Ray in the latter might lead to.

Six-banded Armadillo.

Type specimen in Stockholm Museum, its locality unknown,

But Para specimens agree with it in size, and may be provisionally

accepted as typical (ef. P. Z. 8. 1903, ii. p. 242),

142 MR. OLDFIELD THOMAS ON THE MAMMALS

5. SEPTEMCINCTUS. p. dl. Dasypus (= Tate auct.).

Linnean references: Ameen. Acad. i. p. 281, and Syst. Nat.

(6) p. 6, no. 6; but in the latter we are again referred to

the former. Thus the synonymy may be excluded and

primary importance given to the species described. This

is said to be still in the Upsala Museum.

Seven-banded Armadillo.

Type locality not known.

6. NOVEMCINCTUS. p. 51. Dasypus.

Linnean references: Mus. Ad. Frid. p. 6, and Syst. Nat. (6)

p. 6, no. 7, but in the latter no further detail is added

and we must take the former alone.

Nine-banded Armadillo.

Prof. Einar Lonnberg informs me that the type specimen is still

preserved in the Stockholm Museum, and this will need expert

examination whenever an attempt is made to work out the

present difficult group.

Genus 18. Errnaceus. p. 52.

Monotypic. Type, #. ewropeus.

Species :

EUROPEUS. p. 02. Erinaceus.

Linnean references to Fauna Suecica, Syst. Nat. (6), and

Iter Goth.

Common Hedgehog.

Type locality (from the Gothlandska Resa, p. 264): Wamlingbo,

S. Gothland Id.

Genus 19. Tatra. p. 52.

Two species. Type by tautonymy: 7’. ewropea (“ Talpa, Gesn.”).

Species :

1. EUROPHA. p. 52. Talpa.

Linnean references to Syst. Nat. (6), Fauna Suecica, and

Iter Scanicum.

Common Mole.

Type locality: Upsala *.

2, ASIATICA. p. 53. Chrysochloris.

Sole reference : Seba, Mus. 1. p. 51, pl. xxxii. figs. 4, 5.

Cape Golden Mole.

Type specimen in British Museum (Lidth de Jeude Coll.),

B.M. No. 67,.4.12.564 (see P. Z.8. 1892, p. 316).

* The ‘ Skanska Resa’ being later in date than the ‘Fauna Suecica,’ one cannot

justifiably fix a type locality from it when the ordmary Upsalan locality is available

from the latter work.

OF THE TENTH EDITION OF LINNAUS., 143

Genus 20. Sorex. p. 53.

Three species. Type by tautonymy: S. araneus (“ Sorex, Faun.

Suec.”).

Species :

1. ARANEUS. p. 93, Sores.

Linnean references: Faun. Suec. no, 33; Syst. Nat. (6) p. 10.

Common Shrew.

Type locality: Upsala.

2. CRISTATUS. p. 53. Condylura.

Sole reference: P. Kalm.

Type locality : Pennsylvania.

3. AQUATICUS. p. 93. Scalops.

Reference to Seba (pl. xxxii. fig. 3), whose figure is of Talpa

europea, but the description is clearly that of an American

Mole, and was no doubt received from Kalm, who is quoted

as authority for the locality.

Jommon American Mole.

Type locality : Philadelphia.

Genus 21. DipELPHis. p. 54.

Five species, Type by selection and elimination: D. marsupialis.

Species :

1. MARSUPIALIS. p. 94. Didelphis.

Linnean references: Ameen, Acad. i. p. 279; Syst. Nat. (6)

p- 10; Mus. Ad. Fr. p. 10.

In the first of these places the primary, and in the others the

sole reference is to Seba, Mus. i. p. 64, pl. xxxix., which should

be taken as the basis of the name.

Suggested type locality : Surinam.

2. PHILANDER. p. 54. Caluromys.

Sole reference: Seba, Mus. i. p. 57, pl. xxxvi. fig. 4.

Philander Opossum.

Type specimen in British Museum, No. 67.4.12.414 (see P.Z.5.

1892, p. 314).

Suggested type locality : Surinam.

3. OPOSSUM. p. 95. Metachirus.

Sole reference: Seba, Mus, i. p. 56, pl. xxxvi. figs, 1-2.

Quica Opossum.

Suggested type locality: Surinam.

144 MR. OLDFIELD THOMAS ON THE MAMMAES

4, MURINA. p. 99. Marmosa.

Linnean reference: Amen. Acad. i. p. 279.—This is an

error, the only Opossum referred to being the large

Didelphis marsupialis.

Second reference: Seba, Mus. i. p. 48, pl. xxxi. figs. 1-2,

which must be taken as the basis of the name.

Murine Opossum.

Co-types in British Museum, Nos, 67.4.12.541-542 (see P. Z.8.

1892, p. 314).

Suggested type locality : Surinam.

5. DORSIGERA. p. 90. Marmosa.

Primary reference to Seba, Mus. i. p. 49, pl. xxxi. fig. 5.

Synonym of Marmosa nvurina.

Type in British Museum, No. 67.4.12.546.

Suggested type locality : Surinam.

Genus 22, Ruryoceros. p. 56.

Two species. Type by tautonymy : R. unicornis (* Rhinoceros,

Jonst.”).

Species :

1. UNICORNIS. p. 56. Rhinoceros.

Linnean reference: Syst. Nat. (6) p. 11, no. 1.

One-horned Indian Rhinoceros.

From Jonston’s ‘Quadrupeds’ (p. 67, 1657), the first work

quoted in the 10th edition, we get the

Type locality: Bengal.

2. BICORNIS. p. 56. Diceros.

Linnean reference: Syst. Nat. (6) p. 11, no, 2, where no

further quotation is given.

Always accepted as the African Two-horned Rhinoceros,

Type locality not determinable, except that the Cape may

generally be considered the place whence early specimens were

brought. “ Habitat in India” may, as usual, be entirely ignored.

Genus 23. Hysrrix. p. 56.

Five species. Type by tautonymy: Z. cristata (“* Hystrix, Gesn.”).

Species :

1. CRISTATA. p. 56. Hystria.

Linnean reference: Syst. Nat. (6) p. 9, no. 1, where the first

quotation is to Ray (Quadr. p. 206), whence we find

Type locality: Rome.

Common Porcupine.

OF THE TENTH EDITION OF LINNAUS. 145

2. PREHENSILIS. p. 97. Coendou.

Linnean reference: Syst. Nat. (6) p.9, no. 2, where the only

quotation is to Ray (Quadr. p. 208). There we get an

account taken from Marcgvave, whence

Type locality: Pernambuco,

Brazilian Percupine.

3. DORSATA. p. 57. Brethizon.

Sole reference: Edw. Aves, p. 52, pl. lil.

Oanadian Porcupine.

Type locality : Hudson’s Bay.

4. MACROURA. p. 07. Atherurus.

Linnean reference: Syst. Nat. (6) p. 2, no. 3, where the sole

quotation is Seba, Thes. 1. p. 84, pl. lit, fig. 1.

Common Brush-tailed Porcupine.

No type locality available.

5. BRACHYURA. p. 07. . Acanthion.

Linnean reference: Syst. Nat. (6) p. 9, no. 4, where the sole

quotation is Seba, Thes. i. p. 81, pl. li. fig. 1,

Malay Porcupine.

Type locality : Malacca.

See Lyon, P. U.S. Nat. Mus. xxxii. p. 579, 1907.

Genus 24. Lepus. p. 57.

Four species. Type by tautonymy: ZL. timidus (‘ Lepus, Gesn.”),

Species :

1. TIMIDUS. p. 57. Lepus.

Linnean references: Syst. Nat. (6) p. 9, no. 2; Faun. Suec.

no. 19.

Variable Hare.

Type locality : Sweden (Upsala).

2. CUNICULUS. p. 58. Oryctolagus.

Linnean references: Syst. Nat. (6) p. 9, no. 3; Faun. Suec.

no. 20.

In the former a reference is again given to the latter.

Clearly based on the Domestic Rabbit (‘ pupillis rubris”).

Later on Linneus became acquainted with the Wild Rabbit,

and stated in the 10th edition :—

“ Habitat in Europa australi.”

3. CAPENSIS. p. 08. Lepus.

Common Cape Hare.

Type locality : Cap2 of Good Hope.

T have considered the question as to whether this might not be

Proc. Zoou. Soc.—1911, No. X. 10

146 MR. OLDFIELD THOMAS ON THE MAMMALS

a Rooi-haas (Pronolagus), but think the length of the tail too

great, while in any case the evidence is insufficient to upset the

usually accepted determination.

4, BRASILIENSIS. p. 58. Sylvilagus.

Linnean reference: Syst. Nat. (6) p. 9, no. 1, where the sole

quotation is Ray, p. 205, whose account is taken from

Maregrave.

Hence

Type locality : Pernambuco.

Genus 25. Castor. p. 58.

Two species. Type by tautonymy: C. fiber (“ Castor, Gesn.”).

Species :

1. FIBER. p. 08. Castor.

Linnean references: Faun. Suec. no. 23; Syst. Nat. (6) p. 10,

MO. Sle

Tn the latter the only reference is to the former, in which we find

Type locality: Lapland.

2. MOSCHATUS. p. 59. Desmana.

Linnean references: Faun. Suec. no. 24; Syst. Nat. (6) p. 10,

no. 2.

Type locality: Russia.

Genus 26. Mus. p. 59.

Sixteen species, Type by tautonymy: M. musculus (‘“ Mus,

Gesn.”).

Species :

1, PORCELLUS. p. 59. Cavia.

Linnean references: Westgoth. Resa, p. 244 (misprinted

224); Syst. Nat. (6) p. 10, no. 1.

Domesticated Cavy.

2. LEPORINUS. p. 59. Dasyprocia.

Sole reference: Catesby, Carol. 11. (7. e. u. Appendix) p. 18,

pl. xvii.

An indeterminable species of Dasyprocta. Tail too short and

size too large for Myoprocta.

No type locality available.

3. LEMMUS. p. 59. Lemmus.

Linnean references to Fauna Suecica and Syst. Nat. (6), as

well as to Act. Stockh. 1740, p. 320, pl. vi. figs. 4 & 5,

where Linnzus gives an account of the Norway Lemming.

Type locality: Lapland.

OF THE TENTH EDITION OF LINN AUS. 147

4. MARMOTA. p. 60. Marmota.

Linnean reference: Syst. Nat. (6) p. 10, no. 11, where the

sole quotation is Ray, p. 221.

Alpine Marmot.

Type locality ; Swiss Aips.

5. MONAX. p. 60. Marmota.

Sole reference: Edwards, Aves, p. 104, pl. ev.

W oodchuck,.

Type locality: Maryland.

6. CRICETUS. p. 60. Cricetus.

Linnean reference: Syst. Nat. (6) p. 10, no. 10, where the

sole quotation is Ray, p. 221.

“Tn Turingia and Misnia reperitur”—consequently

Type locality : Thuringen.

Common Hamster.

7. TERRESTRIS. p. 61. Arvicola.

Linnean references: Faun. Suec. no. 29; Syst. Nat. (6)

DaOS mod

Swedish Water- Vole.

Type locality: Upsala.

8. AMPHIBIUS. p. 61. Arvicola.

Primary reference: Ray, Quadr. p. 217.

Ray’s account is quoted from Willughby, and, as both were

Knglishmen, the animal should be taken as the einen Water-

Vole.

Type locality: England.

9. RATIUS. p. 61. Vpumys.

Linnean references: Faun. Suec. no. 28; Syst. Nat. (6) p. 10,

no, 6.

Black Rat.

Type locality: Sweden (Upsala).

10. MuscuLus. p. 62. Mus.

Linnean references: Faun, Suec. no. 31; Syst. Nat. (6) p. 10,

no, 8.

House- Mouse.

Type locality : Sweden (Upsala).

1]. AVELLANARIUS. p. 62, Muscardinus.

Linnean reference: Faun. Suec,. no. 32.

Dormouse,

Type locality : Sweden (Upsala).

10m

148 MR. OLDFIELD THOMAS ON THE MAMMALS

12, syLvatTicus. p. 62. Apodemus.

Linnean references: Faun. Suee. no. 30; Syst. Nat.(6) p. 10,

TOy fe

Long-tailed Field-Mouse.

Type locality : Sweden (Upsala).

13. sTRIATUS. p. 62. Arvicanthis.

Linnean reference: Mus. Ad. Frid. p. 10.

The punctated white lines described indicate that this mouse

is the species usually known as Arvicanthis pulchellus Gray.

-Having been brought in a ship coming from ‘“ India,” its locality

is no doubt that part of W. Africa where such a ship would have

touched. Consequently we may consider its

Type locality: Sierra Leone.

14. LONGIPES. p. 62. Alluctaga.

Linnean reference: Mus. Ad. Frid. p. 9.

A five-toed Jerboa, not specifically determinable without

examination of the type, if one exists.

15. sacuLus. p. 63. Jaculus.

References: Linn. in Hasselq. Itin. p. 198; Act. Stockh.

1752, p. 123.

Lesser Egyptian Jerboa.

Type locality: Lower Egypt.

16. VOLANS. p. 63. Scvuropterus.

Primary reference: Ray, Quadr. p. 215, whence we get

Type locality : Virginia.

American Elying-Squirrel.

Other references are given to Seba, who figures a young

Petaurista, and Edwards, whose animal was drawn from a living

specimen brought from N. America.

Genus 27. Scrurus. p. 63.

Seven species. Type by tautonymy: S. vulgaris (‘‘Sciurus,

Gesn.”).

Species :

1. VULGARIS. p. 63. Sciurus.

Linnean references: Faun. Suec. no. 21; Syst. Nat. (6) p. 9,

Ws, Ihe

Common Squirrel,

‘Type locality : Sweden (Upsala).

OF THE TENTH EDITION OF LINN US. 149

2. NIGER. p. 64.

Sole reference: Catesby, Carolina, 1. pl. lxxiii.

Southern Fox-Squirrel.

Type locality: Carolina. Considered by Bangs to be Southern

South Carolina.

See Bangs, P. Biol. Soc. Wash. x. p. 147, 1896.

3. CINEREUS. p. 64. Sciurus.

References: Ray, Quadr. p. 215; Catesby, Carolina, 11. p. 74,

pl. Ixxiv.; Kalm. Itin. 11. p. 409.

Ray’s account 1s short and unimportant, and Catesby’s is clearly

the primary one. Mr. Bangs has allocated Catesby’s animal to

the Southern Fox-Squirrel (Sczwrus niger) in his work on Eastern

N.-American Squirrels, and his determination may be accepted.

Type locality: coast region of Carolina.

4. FLAYUS. p. 64.

Linnean and sole reference: Ameen. Acad. i. p. 281.

“ Habitat in America.”

Indeterminable, except by reference to the type, if one exists.

In the 12th edition, Carthagena is added as the locality, but

on what grounds does not appear.

5. GETULUS. p. 64. Atlantoxerus.

References: Ray, Quadr. p. 216; Edwards, Aves, pl. 198;

Seba, Mus. i. p. 76, pl. xlvii. fig. 3.

The first two both refer to the Barbary Squirrel, and the third,

which is a Chipmunk, may be ignored.

Type locality : Barbary.

6. SERIATUS. p. 64. Tamas.

Linnean reference: Mus. Ad. Frid. p. 8, where a quotation

is given of Catesby, Carolina, ii. pl. Ixxv.

Common Eastern Chipmunk.

Type locality : ‘‘ Virginia and Carolina.”

7. VOLANS. p. 64. Sciuropterus.

Linnean references: Faun. Suec. no. 22; Syst. Nat. (6) p. 9.

Northern Flying Squirrel, Sciwropterus russicus, the name

volans being already used (from Mus voluns L.) in the genus.

Type locality: Finland.

Genus 28. CamEuus. p. 65.

Four species, of which two, C. dromedarius and bactrianus;

have “ Camelus ” among their synonyms.

Type by tautonymy and elimination: C. bactrianus *.

* Cf. © Opinions of Nomenclature Commissioners,’ 1910, p. 37.

150 MR. OLDFIELD THOMAS ON THE MAMMALS

Species :

1. DROMEDARIUS. p. 65. Camelus.

Linnean reference: Syst. Nat. (6) p. 18, no. 1, where the

sole quotation is of Ray, p. 143.

One-humped Camel.

Type locality : “deserts of Libya and Arabia” (Ray).

2. BACTRIANUS. p. 65. Camelus.

Linnean reference: Syst. Nat. (6) p. 13, no. 2.

Two-humped Camel.

Type locality: ‘ Bactria.”

3. GLAMA. p. 65. Lama.

Linnean reference: Svst. Nat. (6) p. 13, no. 3, where the

sole quotation is of Ray, p. 145.

Llama.

Type locality : Peru.

4. pAcos. p. 66. Lama.

Linnean reference: Syst. Nat. (6) p. 13, no. 4, where Ray

(p. 147) is again alone quoted.

Alpaca.

Type locality: Peru.

Genus 29. Moscuts. p. 66.

Monotypic.

Species :

1. MOSCHIFERUS. Moschus.

Linnean reference: Syst. Nat. (6) p. 13, where Ray is alone

quoted.

Type locality (from Linneus): ‘* Tataria versus Chinam.”

Genus 30, Cervus. p. 66.

Eight species, Type by tautonymy: C. elaphus (*‘* Cervus,

Gesn”’).

Species :

1. CAMELOPARDALIS. Giraffa.

Linnean reference: Syst. Nat. (6) p. 18, no. 1, where

references are given to Bellonius (Obs. ii. p. 119) and Ray,

the latter of whom merely quotes the former.

Giraffe.

Type locality : Egypt (seen in captivity at Cairo).

OF THE TENTH EDITION OF LINNAEUS. 151

2. ALCES. p. 66. Alces.

Linnean references: Syst. Nat. (6) p. 18, no. 2; Faun. Suec.

MOe di.

Elk.

‘Type locality : Sweden.

3. ELAPHUS. p. 67. Cervus.

Linnean references: Faun. Suec. no. 38; Syst. Nat. (6)

De lo, Hoy 3

Red Deer.

Type locality : Sweden.

4, TARANDUS. p. 67. Rangvfer.

Linnean references: Faun. Suec. no. 393; Syst. Nat. (6)

p. 13, no. 4.

Reindeer.

Type locality ; Lapland.

5. DAMA. p. 67. Cervus.

Linnean references: Faun. Suec. no. 40; Syst. Nat. (6) p. 13,

no. d.

Fallow Deer.

** Habitat in vivariis Regis & Magnatum.”

6. BEZOARTICUS. p. 67, Blastocerus.

References :

Mazama, Hern. mex. p. 324.

Cuguacu-ete, Maregr. Bras. p. 235; Piso, Bras. p. 98; Ray,

Quadr. p. 90.

Linnean diagnosis taken from Marcgrave, whose account should

therefore be selected as the primary reference.

Pampas Deer (Cariacus campestris auctorum).

Type locality : Pernambuco.

See Lydekker, ‘ Deer of all Lands,’ p, 287.

7. CAPREOLUS. p. 68. Capreolus.

Linnean references: Faun, Suec. no. 41; Syst. Nat. (6)

p» 13, no. 6.

Roe- Deer.

Type locality : Sweden.

8. GUINEENSIS. p. 68. (2)

Linnean reference: Mus. Ad, Frid. i, Wee

Hab. Guinea.

Not as yet identified,

152 MR. OLDFIELD THOMAS ON THE MAMMALS

Genus 31. Capra. p. 68.

Twelve species. Type by tautonymy: C. hircus (“‘ Capra, Gesn.”).

Species :

1. HIRCUS. p. 68. Capra.

Linnean reference: Faun. Suec. no. 42.

Domestic Goat.

2, IBEX) es 10G. Capra.

Linnean reference: Syst. Nat. (6) p. 14, no. 6.

Reference to Ray only—p. 77 (misprinted 79).

Swiss Ibex.

Type locality : Swiss Alps— Valais.

3. RUPICAPRA, p. 68. Rupicapra.

Linnean reference: Syst. Nat. (6) p. 14, no. 5.

Chamois.

Type locality : Swiss Alps.

4. DEPRESSA. p. 69. (2)

Linnean reference: Syst. Nat. (6) p. 14, no. 2, where a further

reference is given to the 3rd and 4th editions of the

‘Systema,’ but no additional information is there given.

Indeterminable.

5. REVERSA. p. 69. (2)

Linnean reference: Syst. Nat. (6) p. 14, no. 3, where also

the 3rd and 4th editions are quoted.

Indeterminable.

6. PYGMEA. p. 69. Neotraqus.

Linnean reference: Syst. Nat. (6) p. 14, no. 4, where Seba

Gi. p. 70, pl. xliii. fig. 3) is alone quoted.

Royal Antelope.

Type locality: Guinea.

7. GAZELLA. p. 67. Oryx.

Linnean reference: Syst. Nat. (6) p. 14, no. 8, where Ray’s

Gazella indica (p. 79) is alone quoted.

Common Gemsbok. |

Locality not defined, but may be taken as 8. Africa.

8. CERVICAPRA. p. 69. Antilope.

Linnean reference: Syst. Nat. (6) p. 14, no. 7, where there

are two quotations—

Gazella africana, Ray, Quadr. p. 79.

Capricerva, Kempf. Ameen. p. 398, pl. 401, fig. 1.

Both refer to the Indian Black-Buck.

Type locality : India.

OF THE TENTH EDITION OF LINN AUS. 153

9. DORCAS. p. 69. Gazella.

Linnean reference: (6) p. 14, no. 9, where the sole quotation

is Ray, p. 80. ;

Common Gazelle.

No definite locality.

10. arimmta. p. 70. Cephalophus.

Linnean reference: Syst. Nat. (6) p. 14, no. 10, where the

only quotation is to Ray’s Capra sylvestris africana

grimmit, which again refers to “ Ephem. German. An. 14,

Obs. 57,” ¢.e. Grimm, Misc. cur. Acad. Nat. Cur., Decas ii.

Ann. iv. p. 131 (1686).

Common Duiker.

Type locality : Cape Town.

11. mamBrica. p. 70. Capra.

Linnean reference: Syst. Nat. (6) p. 14 (mo. 11), where the

primary quotation is to Ray, p. 81.

A long eared domestic Goat.

Type locality: Syria.

12. ammon. p. 70. Ovis.

Sole reference: Gesn. Quadr. p. 155, but the diagnosis and

‘Habitat in Siberia, Gmelin,” indicate another and more

essential source of information, which may be traced to

J. G. Gmelin, Reise durch Siberien, i. p. 368 (footnote),

Gene

Siberian Wild-Sheep.

Type locality: Upper Irtisch R., Siberia.

Genus 32. Ovis. p. 70.

Three species. Type by tautonymy: O. aries (“ Ovis, Gesn.”).

Species :

Ie -Arimss p70! Ovis.

Linnean references: Faun. Suec. no. 43; Syst. Nat. (6)

pals, no: 1.

Domestic Sheep.

Type locality : Sweden.

2. GUINEENSIS. p. 71. Ovis.

Linnean reference: Syst. Nat. (6) p. 15, no. 3, where the

reference is to Ray, who in turn quotes Marcgrave.

Domestic Sheep of Guinea.

3. STREPSICEROS. p. 71. Ovis.

Linnean reference: Syst. Nat. (6) p. 15, no. 2, whence

Bellonius, Obs. i. p. 20, 1605.

Cretan Domestic Sheep.

i oye! MR. OLDFIELD THOMAS ON THE MAMMALS

Genus 33. Bos. p. 71.

ive species. Type by tautonymy: B. taurus (* Bos, Gesn.”).

Species :

LATS Syn ose Bos,

Linnean references: Faun. Suec. no. 44; Syst. Nat. (6)

pel omnomle

Domestic Ox.

Type locality : Sweden (Upsala).

2. BONASUS. p. 71. Bison.

Linnean reference: Syst. Nat. (6) p. 15, no. 2, where there

is no further reference, so we may take that in the 10th

edition to the Bonasus of Ray (p. 71).

European Bison.

No type locality available.

3. BISON. p. 72. Bison.

Linnean reference: Syst. Nat. (6) p. 15, no. 3, where the

sole quotation is the ‘* Zaurus mexicanus” of Hernandez,

Mexico, p. 587.

American Bison. .

Type locality: “ Mexico.”

4. BUBALIS. p. 72. Bubalus.

Linnean reference: Syst. Nat. (6) p. 15, no. 4, where a

diagnosis, but neither further reference nor locality, is

given. Passing to the next reference in the 10th edition,

we get Ray (p. 72), whose account is based on the domesti-

cated Buffalo. Therefore

Type locality : Italy (Rome).

5. INDICUS. p. 72. Bos.

Sole reference: Edwards, Aves, iv. p. 200 (1751).

Indian Humped Ox,

Type locality : E. Indies.

Genus 34, Equus. p. 73.

Three species. Type by tautonymy: £. caballus (** Equus,

Gesn.”).

Species :

]. CABALLUS. p. 73. Hquus.

Linnean reference} Faun. Suec. no, 84,

Domestic Horse,

OF THE TENTH EDITION OF LINNAUS. 55

2. ASINUS. p. 73. Equus.

Linnean reference: Faun. Suec. no. 35.

Domestic Ass.

3. ZEBRA. p. 74. Equus.

Linnean reference: Syst. Nat. (6) p. 11, no. 3, where a

diagnosis, but no locality or further reference, is given.

Passing to the next reference we have Edwards, Aves, v.

pp- 27 & 29, pls. 222 & 223, the first of which is a Mountain

Zebra, and the second a Quagga. The first should be

taken as typical.

Type locality: Cape of Good Hope.

Genus 35. Hirrororamus. p. 74.

Two species. Type by tautonymy: H. amphibius (‘+ Hippo-

potamus, Bell.”).

Species :

1, aAMpHIBIUs. p. 74. Hippopotamus.

No Linnean reference, though the species occurs Syst. Nat.

(6) Sprite

Hippopotamus.

Type locality: R. Nile.

2. TERRESTRIS. p. 74. Tapirus.

Two references: Tapiierete, Maregr. Bras. p. 229; Ray,

Quadr. p. 126, where Maregraye is again quoted.

Brazilian Tapir.

Type locality: Pernambuco.

Genus 36. Monopoy. p. 75.

Monotypic.

Species :

1. MONOCEROs. p. 75. Monodon.

Linnean references: Faun. Suec. no. 263; Syst. Nat. (6) p. 39.

Narwhal.

‘“ Habitat in mari atlantico.”

Genus 37. Bauz@ya. p. 75,

Four species. Type by tautonymy: B. mysticetus (“ Balena,

Val).

Species :

1. MYSTICETUS. p. 75. Baleena.

Linnean references: Faun. Suec. no. 264; Syst. Nat. (6)

p. 39, no. 1,

In these places the species 1s called ‘*Gronlands Waltisk” and

156 MR. OLDFIELD THOMAS ON THE MAMMALS

* Gronlandswal ” respectively, so that the Greenland species of

Right Whale may be at once accepted.

Type locality ; Greenland seas.

Mr. True, in his ‘ Nomenclature of the Whalebone Whales’ *,

comes to the same conclusion, for the same reason, after two

pages of discussion of what I venture to consider negligible

synonyms.

2. PHYSALUS. p. 75. Balenoptera.

Linnean references: Faun. Suec. no. 265; Syst. Nat. (6)

Toe OM), OK 2s

In both these places the reference is to Artedi (Gen. 77,

Syn. 107), to which alone validity should be attached.

On reference to Artedi, we find Ray (Syn. Pisce. 9) quoted

primarily, who in turn gives an abbreviated account of Martens’s

‘* Finfisch.” This has been shown by Mr. True to be a Common

Rorqual, to which, in agreement with him, I think the name

physalus should be applied.

Type locality : Spitzbergen seas.

The result is therefore the same as Mr. True’s, but the steps are

formal, strict, and inevitable, and do not involve any balancing

of the importance to be attached to different synonyms.

3. Boops. p. 76. Balenoptera.

Linnean reference: Mus. Ad. Frid. i. p. 51 (misprinted 50).

But this reference is wrong as to page, and the diagnosis is a

repetition of that of the succeeding species, B. musculus. It may

therefore be ignored.

The next reference is to species iii. on p. 107 of Artedi’s

‘Synonyms, where in turn Ray’s “ Balena tripinnis nares

habens ....” is quoted. The latter is based on Sibbald’s

account of a whale which Mr. True shows to be the same as

the last species, B. physalus, of which, therefore, b. boops is a

synonym.

Type locality: Firth of Forth, Scotland.

4. MUSCULUS. p. 76. Balenoptera.

Linnean reference: Syst. Nat. (6) p. 39, no. 3.

Here we get Artedi, Syn. p. 107, whence come references to

Ray (Pisce. p. 17), and, at last, the real basis of all, Sibbald’s

‘ Phalainologia.’

Sibbald’s Rorqual.

Type locality: Firth of Forth, Scotland.

* P.U.S, Nat. Mus. xxi. p. 617, 1898.

fee ee ee ee

Lt. ee ee

AAT lal ni etal tn i 8 <

OF THE TENTH EDITION OF LINN EUS. 157

Genus 38. Puyserer. p. 76.

Four species nominally, but all prove to be the Sperm Whale,

so that the genus is really monotypic.

Species :

1. caropon. p. 76. Physeter.

Linnean reference: Syst. Nat. p. 39, no. 1 (of Catodon, not

Physeter), where there is a reference to the ‘‘ Catodon

fistula in rostro” of Artedi(Syn. p. 108), by whom in turn

Ray and Sibbald are quoted.

Ray’s account is abbreviated from that of Sibbald, whose

Balena minor *, from Kairston, Orkneys, is the real basis of the

name.

This animal has been said by some authors to be the Beluga,

and by others the Sperm Whale, and after a careful consideration

of the question, with the kind and highly opportune assistance

of Dr. Einar Lonnberg, I am decidedly of opinion that it was the

latter species.

The absence of teeth in the upper jaw is a definite character, to

which much weight should be attached ; while the fact that females

of the Sperm Whale go together in schools would account for the

large number (105) stranded at Kairston, and their comparatively

small size (24 feet).

As a consequence, the Sperm Whale should bear the name

of Physeter catodon L., of which the next name would be a

synonym.

Type locality : Kairston, Orkneys.

2. MACROCEPHALUS. p. 76. Physeter.

Linnean references: Faun. Suec. no. 262; Syst. Nat. (6)

p. 39, no. 2 (of Catodon).

Sperm Whale (Physeter catodon, see above).

Type locality : Norwegian seas.

3. MICROPS. p. 76. (Physeter.)

Linnean reference: Syst. Nat. (6) p. 39, no. 1 (of Physeter),

where Artedi (Syn. 104, Gen. 74) is alone quoted.

MEP GURSTO: | PoNit. (Physeter)

Linnean reference: Syst. Nat. (6) p. 39, no. 2 (of Physeter),

where Artedi (Syn. 104, Gen. 74) is alone quoted.

These two names refer to the mythical ‘‘ High-finned Cachalot,”

whose origin no doubt lies in faulty observation of ordinary

Cachalots, and they may be considered as synonyms of Physeter

catodon,

158 MR. OLDFIELD THOMAS ON

Genus 39. DELPHINUS. p. 77.

Three species. Type by tautonymy: D. delphis (‘‘ Delphinus,

Bell.”).

Species :

1. PHOCENA. p. 77. Phocena.

Linnean references: Faun. Suec. no. 266; Syst. Nat. (6)

jee @a IO, Il

Common Porpoise.

Type locality : Swedish seas.

2. DEUP EIS. pail. Delphinus.

Linnean reference: Syst. Nat. (6) p. 39, no. 2, where Artedi

(Syn. 105) is alone quoted.

Common Dolphin.

Type locality : ‘‘ Huropean seas.”

SMORCA ENED: 0. Orcinus.

Linnean references: Faun. Suec. no. 267; Syst. Nat. (6)

[Dn BA WO, 35

Common Killer.

Type locality : North Sea.

10. The Duke of Bedford’s Zoological Exploration of Haste:n

Asia.— XIII. On Mammals from the Provinces of

Kan-su and Sze-chwan, Western China. By OLDFI“uLD

Tomas, F.R.S., F.Z.S.

[Received and Read February 7, 1911.]

After making the fine collection from Shan-si and Shen-si,

which formed the subject of No, XI. of the present series of

papers, the Duke of Bedford’s collector, Mr. Malcolm Anderson,

paid a visit to Europe and America, and then returned again to

China in the autumn of 1909. He there obtained the services of

Dr. J. A.C. Smith and Mr. Kingdon Ward, who accompanied

him in his further explorations inland.

The party first went up into Southern Shen-si, whence a small

collection was sent home, which included the three species whose

descriptions were published in the ‘ Proceedings’*. ‘The other

forms then obtained will be referred to in a succeeding paper.

The next collection was made in the mountain region of 5.W.

Shen-si, but, owing to accidental delays, has reached London

later than the succeeding set, which forms the subject of the

present paper.

Mr. Anderson’s party then proceeded into Kan-su, a province

* P.7.S. 1910, p. 635. These descriptions, though not so numbered, may be

considered to form No, XTI. of the present series of papers.

MAMMALS FROM WESTERN CHINA, 159

hitherto entirely unrepresented in our National Museum. The

collection there made and now described is of the utmost interest

and greatly increases our knowledge of the Chinese Fauna.

Then a move was made southward into Western Sze-chwan,

the main objective of Mi. Anderson’s long journey. For it was

in this region that the famous collections of Pére David were

formed, and authentic specimens of the many species he dis-

covered were vital desiderata for all satisfactory work on Chinese

Mammals. Collections were made at and near Ta-tsien-lu, and

again with great success on the sacred mountain Omi-san, near

Kia-ting-fu.

The result of the party’s labours forms, I believe, the finest

collection of small mammals that has ever come from China. No

less than 16 species and subspecies have had to be described as

new*, while authentic specimens have been obtained of nearly

all Pere David’s discoveries.

Special mention may be made of the series of .Veotetracus, a

peculiar Insectivore recently described by Prof. Trouessart, of the

black- striped Shrews, no such coloration having been previously

described in the Family, of the specimens of U ‘ropsilus and

Blarinella, and of the oneness Vole P ‘oedromys bedfordi, the

type of a new genus. In all there are 360 specimens, belonging

to 48 species and subspecies.

The collection, as a whole, gives striking evidence of the uni-

formity of the mammal fauna of China north of an east and west

line at about 34° N., corresponding in its western part with the

T'sin-ling range of mountains. Specimens from Korea on the

east, through Shantung, Shan-si and Shen-si to Kan-su on the

west, are all remarkably uniform in character, while there is

an abrupt change on passing from Kan-su to the southern side

of the range into Sze-chwan. The inhabitants of the former are

all ‘‘desert-coloured” animals, and of the latter dark animals—

no doubt because of the more forested nature of the country.

Zovlogists have every reason to be grateful to the Society’s

President for this fine addition to the available material for

working out the Hastern Asiatic Fauna. It forms a worthy

supplement to the results that have been already attained by the

same means in Japan, Korea, and other parts of China, the whole

constituting one of the most ‘magnificent series of collections that

has ever been obtained.

Considering again the hardships that had to be undergone, the

roughness and ald nature of the country, the Taiiendiiness of

the Uones. at whose hands Mr. J. W. Brooke had recently met

his death, and the severe climatic conditions, we must extend our

admiration to Messrs. Anderson, Smith, and Ward, for the

extent of the collection and the excellent Eondition in which they

have succeeded in sending it home.

* The complete-account of these new forms appears in this communication, but

since the names and preliminary digenoses were published in the ‘ Abstract,’ No. 90,

1911, these species are distinguished by the names being underlined.—Epsror.

160 MR. OLDFIELD THOMAS ON

|]. RHINOLOPHUS CoRNUTUS PUMILUS K. And.

6. 2560, 2561, 2563. @. 2562, 2564. Kia-ting-fu, 8. Sze-

chwan. 1200’.

Apparently indistinguishable from the type, with which

Dr. Andersen has compared it.

2. BARBASTELLA DARJELINGENSIS Hodgs.

6. 2523.) Omiesaneoo 00.

A specimen of this rare species was also obtained in Sze-chwan

—at Yin-shin-wan, N.W. of Chong-tu—by Mr. W. N. Fergusson.

T can find no difference between these specimens and Hodgson’s

type from Darjiling.

3. PLECOTUS ARIEL.

Thos. Abstr. P. Z.S. 1911, p. 3 (Feb. 14).

@. 2343. Ta-tsien-lu, Sze-chwan. 8400!. B.M. No. 11.2.1.6.

Collected 23 Jane, 1910. Type.

Most nearly related to P. wardi, but darker.

Size among the largest of the genus. Fur of medium length.

General colour dark “ broccoli- brown,” the ends of the hairs dull

dyvab. Under surface ‘paler drab, the bases of the hairs dark

slaty. Muzzle blackish. Proportions of thumbs and hind limbs

as in sacrimontis.

Skull large, with swollen brain-case and broad interorbital

region. Bulle large, only less than those of the Egyptian species

P£. christiet.

Dimensions of the type, the starred measurements taken in the

flesh :—

Forearm 44 mm.

Head and body 53* mm,; tail 57*; ear 43*; third finger,

metacarpus 39°5, first phalanx 15-5; thumb (c. u., exclusive of

metacarpal) 9°2 ; lower leg and hind foot (c. u.) 31.

Skull: greatest length 17-2; basi-sinual length 13-2; zygomatic

breadth 9; intertemporal constriction 4; mastoid breadth 9-4;

front of canine to back of m* 5°6; greatest horizontal diameter

of bulle 4-6.

Hab. and Type as above.

Of the genus Plecotus, P. homochrous (Nepal) and puck (Murree)

(doubtfully distinct from each other) stand aside from the rest

owing to their narrow brain-case ; auritus (Kurope) has rounder

skull, small bulle, and short thumbs; christier* (Egypt) large

brain-case and very large bulle; wardi (Ladak and Kashmir)

large skull, rather large bulle, long thumbs, and very pale colour ;

sacrimontis (Japan) large skull, rather small bulle, and long

thumbs; and finally the present species has large rounded skull,

large bulle, long thumbs, and dark colour. No doubt it is most

* Although the name egyptiacus is quoted both hy Dobson and Trouessart from

Is. Geoffroy’s paper of 1882, I fail to find anything but “ Oreillard d’Egypte ” in

that paper, and therefore use Gray’s name of 1838,

MAMMALS FROM WESTERN CHINA. 161

nearly allied to P. wardi, but the colours of the two are nearly at

the opposite ends of the scale.

4. Myotis ALTARIUM.

Thos. Abstr. P. Z.S. 1911, p. 3 (Feb. 14).

6. 2427, 2428, 2429, 2431, 2433. 9. 2422-2496, 2430, 2432.

Omi-san, Sze-chwan. 6000’.

A large species—about the size of JZ. bechsteini—the skull

remarkably shortened.

Size large as compared to ordinary small species of d/yotis.

Fur rather long, but thin and poor, at least in summer pelage;

hairs of back about 8 mm. in length. General colour above

uniform dull brown (paler than “ Prout’s brown ”), the tips of the

hairs rather lighter. Under surface little lighter than upper,

except that the tips of the hairs are more distinctly lighter. Hars

long, nearly as long as in MW. bechsteini, but rather narrow; inner

edge evenly convex, outer slightly concave above, convex in lower

half, a strongly developed antitragal lobule at its base, separated

by a deep notch. Tragus long, not very sharply pointed, evenly

but slightly curved outwards; a well-marked lobule at its outer

base. Membranes naked throughout; no fringe on interfemoral.

Feet rather above normal size, but not as in “‘ Lewconoz”; calcar

extending rather more than halfway towards the tip of the tail;

a very narrow postcalcareal lobule.

Skull different in shape from that of ordinary J/yotis, owing to

the shortening of the rostrum, which is broad and evenly narrowed

forwards, instead of there being a narrow and nearly parallel-sided

anterior portion. This condition is, however, approached in

M. pequinius. Upper outline without frontal convexity, nearly

straight from its highest point near the lambda to a point over

the small premolars, then abruptly concave, the short nasal region

retroussée. Palate unusually vaulted.

Middle upper premolar about half the size of the anterior, both

in the tooth-row; and the same is the case with the lower

premolars,

Dimensions of the type, the starred measurements taken in the

flesh :—

Forearm 45 mim.

Head and body 55* ; tail 48* ; ear 22*; tragus on inner edge 8 ;

third finger, metacarpus 40, lst phalanx 13:3; lower leg and

foot (ec. u.) 29.

Skull: greatest length 15-2, basi-sinual length 12; breadth of

brain-case 7°9; front of canine to back of m? 6:5.

Hab. as above.

Type. Adult female. B.M. No. 11.2.1.9. Original number

2423. Collected 2 August, 1910.

This is a most peculiar species and readily recognizable by its size,

long narrow ears, and the unusual shape of its skull, which differs

considerably from that of most members of the genus, although

another Chinese species, JZ. pequinius, shows an approach to it.

Proc. Zoou. Soc.—1911, No. XI. I

162 MR. OLDFIELD THOMAS ON

5. Myotis MouPINENSIS M.-Edw.

3. 2322. 23 miles §.E. of Ta-tsien-lu, Sze-chwan. 7500’.

Milne-Edwards’s Vespertilio moupinensis was placed by Dobson

as a synonym of V. muricola Hodgs., but is readily distinguished

by the sharp notch in the outer edge of the ear.

Two specimens of it were also obtained by Mr. W. N. Fergusson

at Yin-shin-wan, N.W. of Chong-tu.

6. NEOTETRACUS SINENSIS Trouess.

3. 2416. 45 miles 8.W. of Ya-chow, W. Sze-chwan.

3. 2445, 2459 (yg.), 2460 (yg.), 2467, 2468, 2469, 2471, 2480.

@. 2434 (imm.), 2444, 2446, 2450, 2472. Omi-san. 6000’.

This series of the highly interesting Insectivore, Veotetracus,

recently described by Prof. Trouessart, forms a most valuable

addition to the Museum Collection, and at the same time enables

me to increase our knowledge of its distribution and characteristics.

The chief point I have to note is its milk dentition, Mr. Ander-

son’s series including one half-grown and two quite young

specimens.

The result of my examination is the following formula, showing

both the milk and permanent teeth :—

femiat 7O (1 Ca RON BO

a eG ler | Ones ead Lee alee}

Li§ == @2 5 24S —_ i, ——

jhe Bags fie. | Owes ee eos:

lite aa ae) ul eee oe

Comparing this with the formule of other members of the

group *, we find that it is almost precisely the same as that of

fylomys, agreeing with it even in the absence of a third upper

milk incisor, and the presence of the corresponding tooth below.

The main difference is in the entire absence, both above and

below, in milk and permanent series, of p*, this tooth being present

in both milk and permanent series of Hylomys above, and in the

permanent series below.

The proportions of the milk-teeth are not very different from

those of Hylomys, though di’ surpasses di° to a greater extent.

While the permanent upper carnassial (p*) is of the usual shape

characteristic of the family, its predecessor (dp*) is quite peculiar

in the entire suppression of the small postero-internal cusp and

the reduction of the antero-internal, these being in Hylomys

well developed and approximately similar in position to those on

the permanent p*. ‘They are also both present in Gymnra and

Hrinaceus.

Taking all characters into consideration, I am of opinion that

Neotetracus is most nearly related to Hylomys and is strictly a

member of the Gymnurine, without any special leaning towards

the Erinaceine.

* Of. P.Z.S. 1892, p. 505.

MAMMALS FROM WESTERN CHINA, 163

“In damp forest; no water.

“Stomach contents, earthworms.

“4 embryos—mamme 2—2= 8, another specimen with

5 embryos.”—M. P. A.

7. Tapa Lonerrostris M.-Edw.

Gg. 2538. Omi-san. 9500’.

An old individual from Yung-li-pa, N.W. Sze-chwan, obtained

by one of Mr, Styan’s collectors, was previously the only specimen

of this species in the Museum.

8. Urorsitus soricirpes M.-Edw.

6. 2457, 2504, 2520, 2540, 2550, 2554. 9. 2489, 2527

2539. Omi-san. 6000'—9500'.

Although, owing to their being occasionally deciduous, the

teeth of this animal may have been quite correctly indicated for

the type specimen by Prof. Milne-Edwards, the full normal

number of teeth would appear to be greater both above and

below than he stated. For, above, in front of the larger p* there

is generally present a minute tooth smaller than any other in the

jaw, and in the lower jaw there is a similarly minute tooth

immediately following the large anterior incisor. Neither of

these is indicated by Milne-Edwards. The full number of the

permanent teeth would therefore seem to be

1, C.7, P.3, MoS 4X2 = 38,

the same as in Dymecodon. In the milk-dentition, however,

there is an interesting difference, as there are three lower milk-

premolars in addition to the non-changing p.,.

Homologising the individual teeth, the full formula would

therefore seem to be :—

(lee ors (et fois eee Se"

a ae a | | aoa ie oes

Z C.<— PR. <—— ow ee

} 0.2.3 [2 PI BALE ores

(OWS 2 2S et ale? One Ber a

This formula may be compared with that of Dymecodon, as given

in a previous paper of the present series * .

The chief differences in the dentition of the two genera are the

minute size (and occasional absence) of p* in Uropsilus, this tooth

in Dymecodon being larger than p', the rounder, less carnassial

shape of p*in Uropsilus, and, in the lower jaw, the extremely

minute size of the posterior of the two incisors (i,) and a con-

siderable superiority in size of p, over p,, these teeth being

‘subequal in Dymecodon.

This interesting Insectivore had not been previously represented

in the British Museum Collection.

* P.Z.S 1908, p. 51 (footnote).

11*

164 MR. OLDFIELD THOMAS ON

9, Sorex cyLrypricauDA M.-Edw.

3. 2507, 2547. 9. 2548, 2555. Omi-san. 9500’.

These Shrews seemed on arrival to represent a totally new form

of the group, for they are conspicuously marked with a deep black

line running down the centre of the back, a coloration hitherto

unrecorded in the family.

But after a personal examination, kindly facilitated by

_ Prof. Trouessart, of the species described under the above

name by Milne-Edwards, I have come to the conclusion that

Mr. Anderson’s specimens are referable to that animal, although

its describer made no mention of the dorsal line. Probably the

type was skinned out of spirit, and was already discoloured when

he described it.

The skull agrees exactly in size and dental characters with that

of the type.

10. SoREX BEDFORDI.

Thos. Abstr. P.Z.8. 1911, p. 3 (Feb 14).

g. 2323, 2337. 23 miles 8.E. of Ta-tsien-lu, W. Sze-chwan.

3S. 2359, 2374, 2391. ‘Ta-tsien-lu. 9000’.

g. 2441, 2475, 2490, 2506, 2521, 2541. 9. 2496, 2497,

500, 2522, 2524, 2532, 2542, 2549. Omi-san. 6000'—9000’.

A smaller species with a dorsal stripe.

Size decidedly smaller than in S. ecylindricauda, to which it

perhaps bears the relationship that S. mints does to S. araneus.

Fur soft and fine; hairs of back about 4:2-4-4 mm. in length.

General colour above brown, something between “ bistre” and

“ Prout’s brown”; scarcely or not lighter below. Back with an

indistinct dorsal stripe, like that of S. cylindricauda, but usually

less distinct, sometimes only discernible as a faint darker median

wash often broken over the shoulders; in one instance, however

(No. 2532), it is as strongly black as in the larger species. Hands

and feet pale brown. ‘Tail dark brown above and terminally

below, lighter proximally below.

Skull rather smaller than that of S. aranews. Three anterior

unicuspids equal ; two posterior smaller and subequal, the posterior

commonly the larger.

Dimensions of the type :—

Head and body 55 mm.; tail 55; hind foot 13.

Skull: condylo-basal length 17:4, greatest breadth 8-4, upper

tooth-series 7°4 ; front of p* to back of m* 3:5.

Hab. as above. Type from Omi-san, 9500’.

Type. Adult male. B.M. No. 11.2.1.41. Original number

2541. Collected 19 August, 1910.

The characteristic striping of this Shrew is not normally so well

marked as in S. cylindricauda, but it is quite sufficiently evident

in all the specimens to afford a ready means of distinguishing the

species from any of its allies.

I have named this very pretty little species in honour of the

Duchess of Bedford, whose interest in zoology is well known.

MAMMALS FROM WESTERN CHINA. 165

11. SorEX WARDI.

Thos. Abstr. P. Z.8. 1911, p. 3 (Feb. 14).

3. 2230. Tau-chow, Kan-su. 9000’. 25 March, 1910. B.M.

No. 11.2.1.46. Type.

Like S. bedfordic, but paler and with smaller brain-case.

Essential characters as in S. bedfordie. Hairs of back 5°6 mm.

in length. Colour paler, the generai colour of the body nearly

approaching “ drab,” and the tips of the hairs of the under surface

drabby white, markedly contrasting with their slaty bases ; a line

of demarcation perceptible on sides. Dark dorsal streak well

marked, more so than in average S. bedfordie. Hands, feet, and

underside of tail brownish white, the tail much more sharply

bicolor than in bedfordie.

Skull with the brain-case much smaller, and, especially, lower

than in S. bedfordie, its length 4°5 mm. as compared with about

5°D in average specimens of that animal. Unicuspids subequal

throughout, the fourth slightly the smallest. Dental pigment-

ation moderate.

Dimensions of the type :—

Head and body 53 mm.; tail 49; hind foot 12; ear 8.

Skull: condylo-basal length 17, greatest breadth 82; upper

tooth-series 7°4; front of p* to back of m* 3-7.

Hab. and Type as above.

The paler colour, slightly shorter skull-length, and smaller and

much lower brain-case will readily distinguish this Kan-su striped

Shrew from its Sze-chwan ally.

IT have named it in honour of Mr. F. Kingdon Ward, one of

Mr. Anderson’s assistants during the expedition.

12. SoricuLus MAcRuRUS Blanf.

3. 2493. ©. 2503. Omi-san. 9500’.

Agrees closely with Blanford’s type from Darjiling.

13. SoRICULUS SACRATUS.

Thos. Abstr. P. Z.S. 1911, p. 4 (Feb. 14).

3. 2442, 2451, 2476. 9. 2452, 2477, 2485. Omi-san.

6000... °°

A small species allied to S. caudatus ; tail rather shorter than

head and body.

Size smaller than in any described species. Form light and

delicate. Fur fine, velvety; hairs of back about 4:5 mm. in

length. General colour near “slate-grey,” varying in some

specimens towards brown. Under surface paler, glossy “ smoke-

grey,” varying towards “ broccoli-brown.” Hands and feet pale

brown. ‘Tail a little shorter than the head and body, about as in

S. caudatus, brown above, glossy whitish below.

Skull markedly smaller than that of S. caudatus, especially in

the size of the brain-case, the facial part, with the tooth-series,

being nearly as long as in theallied form. Teeth asin S. caudatus;

distinguished from those of the last species by the tooth-rows

166 MR. OLDFIELD THOMAS ON

being further apart anteriorly, so that the median space between

the hinder lobe of the first incisors is about equal in breadth to

that lobe, markedly narrower in S. macrurus.

Dimensions of the type, measured in skin :—

Head and body 60 mm.; tail 54; hind foot (c. u. ) 14.

Skull: greatest length (exclusive of incisors) 181; basal

length U5) 0c greatest breadth 9°5; upper tooth-series 8:1; front

of p’ to back 6 m” 4:4,

Hab. as above.

Type. Adult 2. B.M. No. 11.2.1.54. Original number 2485.

Collected 10 August, 1910.

This is evidently the local representative of S. caudatus, but is

readily distinguishable by its much smaller brain-case.

14, CHODSIGOA SMITHII.

Thos. Abstr. P. Z.S. 1911, p. 4 (Feb. 14).

6. 2849. Ta-tsien-lu. 9000’. 27 June, 1910. B.M. No.

11.2.1.537. Z'ype.

Proportions as in Ch. hypsibia; size much larger.

Size very large, nearly equaling that of the long-tailed Ch.

sdlenskii Kashtch. Fur fine and soft; hairs of back (summer)

3°8 mm. in length. General colour uniform ‘“ mouse-grey” ;

under surface similar in colour, though, owing to the different

texture of the hairs, the surface appears more glossy. Muzzle

scarcely darker than body, Lateral gland not conspicuous, nearer

the hip than the shoulder. Hands and feet brownish white,

lighter terminally, Tail rather shorter than head and body, pale

brown above, whitish below.

Skull much larger than that of Ch. hypsibia, the brain-case

flattened, heavily ridged. Proportions of teeth as in that species,

their tips as usual lightly pigmented.

Dimensions of the type, an old male :—

Head and body 85 mm.; tail 68; hind foot 18.

Skull: condylo-basal length 22° 5, er eatest breadth 10:5; upper

tooth-series 10°2; front of p* to back of m* 5:3.

Hab. and 7 ype as above.

This fine species is conspicuously larger than its ally Ch. hyp-

sibia de Wint., with which I synonymize Ch. berezowski Kashtch,

It is rather smaller than the long-tailed Ch. salenskii Kashtch.

I have named it in honour of Dr. J. A. C. Smith, who ac-

companied Mr, Anderson during the Sze-chwan trip, and assisted

largely in making the collection.

15. BLARINELLA QUADRAYICAUDA M.-Hdw.

6. 2505, 2556, 2558.. 9. 2557. Omi-san. 9500’.

BLARINELLA, &. n.

More allied to the N.-American Slarinw than to any of the

Old World genera of Shrews. Hars very small, though neither

MAMMALS FROM WESTERN CHINA. 167

so rudimentary nor so hair-covered as is the case in Llarina ;

their conch wholly below the surface of the fur. Tail short, but

not nearly so short as in Blarina. Claws large, the anterior

inconspicuously larger than the posterior.

Skull in general build about as in the more delicate species of

the subgenus Crypitotis, though the rostrum is hardly so high.

Teeth numbering 32,as in Sorex and true Blarina. Upper

unicuspids normally 5, but of such proportions and so slanted that

only 3 are as a rule visible in a side view, as in Cryptotis. First

two large and subequal, third about half their size, its hinder

edge level with the front of p'; fourth flattened from before

backwards, about half the size of third, and jammed close against

it, between it and the minute fifth, which is entirely in the inner

half of the tooth-row, close against the large p'. Molars square,

practically without interspaces posteriorly. Lower teeth about

as in Llarina, the large incisors with three notches. All teeth

heavily pigmented, as in Blarina.

Type, B. quadraticauda (Sorex quadraticauda M.-Kdw.)

The discovery that the Llarina group is represented in the Old

World is a most interesting one and very similar to that of the

Sze-chwan Zapus, published by M. Pousargues in 1896. The

present species was described as a Sorex by Milne-Edwards, but

when examining the type in Paris some years ago I felt sure

it was either Llarina or nearly allied to it, and Mr. Anderson’s

series now enables me to place it with certainty.

Of the four specimens obtained three have five unicuspids, as

above described, but in one (No. 2505) the minute fifth is absent

on both sides, so that it seemed possible the type was in the same

condition, only four unicuspids having been described and figured.

A ye-examination of it, which I owe to the kindness of Prof.

Trouessart, shows that it presents the intermediate condition of

having on the right side only 4 unicuspids, the 5th minute one

being missing, just as in no. 2505, while on the left this tooth

is present. Owing, however, to the accidental loss of the 3rd and

4th unicuspids on this left side, the fact that the minute tooth

still present, lying pressed a against p', corresponded to the fifth

tooth that we now know to be normal to the species, had not

been previously observed. There is therefore no doubt whatever

as to the specific identity of the present series with Milne-

Edwards’s animal.

The four groups of short-tailed Shrews may be readily dis-

tinguished from each other by the number and relative visibility

ter)

from outside of the upper unicuspids, as follows :—

5 unicuspids, 4 visible from outside. Dlarina.

5 vs is KA Blarinella.

4 3 Bs se Cryptotis.

5) os 3 " 5 Notiosorex.

So entirely are both posterior unicuspids generally hidden in

Llavinella that the figure of the teeth of LB. (Cryptotis) parva

168 MR. OLDFIBLD THOMAS ON

given by Merriam in his Monograph* would almost do for it,

were it not for the much greater size and posterior extension of

the large upper incisor of the Asiatic animal.

16. CrocrpuRA ATTENUATA M,-Edw.

3. 2279. ©. 2278. Wen-hsien Country, Kan-su.

17. ANOUROSOREX SQUAMIPES M.-Edw.

3g. 2407, 2410, 2412. ©. 2408, 2409, 2411. 45 miles W. of

Ya-chow, Sze-chwan.

6. 2415, 2417, 2420. 2. 2418, 2412. 45 miles S.W. of

Ya-chow.

@. 2435, (24386, 2458, 2464, 2470; QA8l 2485, 2487, 2501;

2533, 2. 2440, 2455, 2465, 2483, 2486, 2488, 2502. Omi-san.

6000'—9500'.

The first-mentioned place would appear to be about 50 miles

S.W. of Moupin, the type locality of the species.

18. VULPES sp.

©. 2201. Near Si-Ho-Hsien, 8. Kan-su.

Apparently representing V. filchneri Matsch. The relationship

of this form to the Moupin Fox (V. hooly David) is not clear.

19. MustreELA RUSSELLIANA.

Thos. Abstr. P. Z.8. 1911, p. 4 (Feb. 14).

6. 2381, 2382. ©. 2388, 2389, 2390. Ta-tsien-lu. 12,000’.

A very small species, with the contrasted body-colours and

tuftless tail of I. kathiah.

Size extremely small, about that of the very smallest known

Weasel—the American M/. rizosa Bangs. Colours of upper and

lower surfaces sharply contrasted, as in J. kathiah and the Stoats,

not as in Jf. nivalis. Upper surface uniform dark brown, rather

less rich and more drabby than in J. kathiah. No darker

markings on face or ears. Under surface a beautiful pinkish

buff, turning into white anteriorly on the chin, interramia, and

lips. Line of demarcation very sharply marked, running from

upper lip to ankle. Arms brown externally, buffy on inner

aspect; hands brown, sometimes mottled with white; hind limbs

similarly brown externally and buffy on the inner aspect; palms

and soles densely hairy. ‘Tail proportionally shorter than in

M. kathiah, slender, not tufted, uniformly brown, the tip not

noticeably darker.

Skull distinguishable from that of all other allied forms by its

extremely small size.

Dimensions of the type :—

Head and body 133 mm.; tail 54; hind foot 22; ear 11.

Skull : condylo-basal length 29:3 ; basal length 27:2; zygomatic

breadth 15:2; interorbital breadth 6-2 ; breadth of brain-case 14 ;

* N,. Am. Fauna, no, 10, pl. iil. fig. 2, 1895.

MAMMALS FROM WESTERN CHINA. 169

palatal length 11; front of canine to back of molar 8 ; p* on outer

edge 2°9.

A male specimen measures: head and body 138 mm. ; tail 54 ;

hind foot 24; carnassial tooth 3:1,

Hab, as above.

Type. Adult female. B.M. No. 11.2.1.86. Original number

2388. Collected 1 July, 1910.

This most beautiful little Weasel competes with the American

M. rizosa Bangs for the honour of being the smallest existing

Carnivore, and is readily distinguishable from any Old World

Mustela by its small size alone. Its essential coloration is very

much as in the much larger MW. kathiak, of which a local race

(dorsalis) has been recorded from this region. But the hind foot

of that animal measures 30 mm., so that it is evidently quite a

distinct species.

The combination Jfustela bedfordi being alveady in use, I have

given to this striking new species a designation based on the

family name of the donor of the present magnificent accession to

the National Collection of Mammalia.

20. TamMiops swINHOEI M.-Edw.

6. 2510. Omi-san, Sze-chwan. 9500’.

21. SctuROTAMIAS DAVIDIANUS M.-Edw.

©. 2277. 40 miles N. of Kai-chow, 8. Kan-su.

22. ScIUROTAMIAS DAVIDIANUS CONSOBRINUS M.-Edw.

6. 2320, 2321. 9.2316. 23 miles S.E. of Ta-tsien-lu. 7500’.

3g. 2456. Omi-san. 6000’.

The Ta-tsien-lu specimens are not very typical consobrinus, but

are none of them adult.

23. MArmota rosusta M.-Edw.

3. 2270. Old Tau-chow, Kan-su. 9000’.

©. 2269. Near Tau-chow. 10,000’.

24. Mus GANSUENSIS Sat.

6. 2220, 2221, 2222. 10 miles 8. of Tau-chow, Kan-su.

Described as a Leggada by Satunin.

25, EPIMYS CONFUCIANUS LUTICOLOR Thos.

3d. 2196, 2197, 2199, 2200. 9. 2198. Near Si-Ho-Hsien,

S. Kan-su. 5000’.

¢. 2206, 2219. 60 miles S.E. of Min-chow, Kan-su. 8000’.

g. 2280, 2286, 2287, 2294, 2295, 2301, 2302, 2309, 2311.

@. 2281, 2282, 2288, 2310, 2312. Wen-hsien Country, S.

Kan-su.

These Kan-su specimens tend to be, as is right, intermediate

170 MR. OLDFIELD THOMAS ON

between the true luéicolor of Northern Shen-si and the darker

confucianus of Sze-chwan. The lighter specimens among them

absolutely match some of the typical series, while the darker ones

match the lighter individuals of the next following set.

26. Hpimys conructAnus M.-Kdw.

6. 2313, 2318. 2. 2314,9315,2317, 2319, 23 miles S:Bet

Ta-tsien-lu, Sze-chwan.

3. 2370, 2371, 2372. Ta-tsien-lu. 9000’.

3. 2405. ©. 2406. 18 miles E. of Ta-tsien-lu. 5500’.

@. 2413. 45 miles W. of Ya-chow.

3. 2443, 2453, 2462, 2463. Omi-san. 6000’.

Pending the arrival of topotypes from Chong-tu, these speci-

mens may be accepted as typical of the true confucianus.

27. Epimys tine Bonh.

@ , 2283, 2289. Wen-hsien Country, Kan-su.

These beautiful ochraceous-buff specimens are quite like the

typical series from Kuatun, and are also very similar to an

example in the Museum from Sikkim, referred—and perhaps

correctly—to #. jerdoni Blyth.

But as this latter determination needs revision with better

material, I use the Chinese name which happens to be

available.

28. HPrIMYS EXCELSIOR.

Thos. Abstr. P. Z.S. 1911, p. 4 (Feb. 14.)

3. 2360, 2379. ©. 2380. Ta-tsien-lu, Sze-chwan. 9000’.

S. 2511, 2526, 2545. Omi-san. 9500’.

General characters of 1. confucianus, but size larger.

Size larger than in confucianus, proportionate length of tail

about the same. Fur long and rather coarse, not mixed with

spines in any of these specimens ; hairs of back about 14 mm. in

length, the isolated bristle-hairs attaining 20. General colour of

adults coarsely lined cinnamon or clay-colour—not unlike bright-

coloured specimens of #. norvegicus. Sides clearer and more buffy.

Belly pure sharply defined white or creamy white, which extends

anteriorly well up over the upper lip. Top of muzzle greyer.

Eyes with darker rims. Ears of medium length, greyish brown.

Hands white above. Feet white with prominent dark metapodial

patches. Tail rather longer than head and body, well-haired and

tufted, brown above for about two-thirds its length, white at the

end and below; hairs at end 7-8 mm. in length.

Skull differing from that of confucianus by its larger size and

more delicate build. Nasals long and slender. Interorbital

region narrow, its borders sharp-edged, but without definite

beads. Brain-case smooth, markedly convex above. Anteorbital

plate but little projected. Palatal foramen long. Bulle small.

Molars decidedly larger than in confucianus.

MAMMALS FROM WESTERN CHINA. tial

Dimensions of the type :—

Head and body 178 mm.; tail 193; hind foot 30; ear 23.

Skull: greatest length 39; basilar length 30°33; zygomatic

breadth 18; nasals 15:2 x 4°3 ; interorbital breadth 4:3 ; breadth of

brain-case 15°6; palatilar length 17; palatal foramina 7:5; upper

molar series 6:8.

Hab. of type. Ta-tsien-lu. 9000’.

Type. Adult male, B.M. No. 11.2.1.131. Original number 2379.

Collected 30 June, 1910.

The true Rats of the genus Hpimys are, as usual, some of the

most difficult of the collection to make out satisfactorily, owing

to their variability in both colour and skull-characters. But it

seems clear that this form from the high ground of Ta-tsien-lu

and the upper part of Omi-san is a distinct species from the

widely spread #. confucianus, differing by its greater size, longer

skull, and, in fully adult specimens, its paler colour—at least, as

compared with the typical dark HL. confucianus of W. Sze-chwan.

It is, of course, a member of the same group of the genus, as is

also the next species, which is again a step larger in size than

LH. excelsior.

29. EPIMYS ANDERSONT.

Thos. Abstr. P. Z.8. 1911, p. 4 (Feb. 14).

2. 2478. Omi-san. 6000’. Collected 8 August, 1910. B.M.

No. 11.2.1.135. Type.

Like ZH. confucianus and excelsior, but very much larger and

with longer tail.

Size large, though not equaling the giant of this group,

Hi. edwardsi. Fur coarse, not spinous ; hairs of back about 10 min.

in length. General colour as in H#. excelsior, a dull-lined clay-

colour, more buffy on the sides. Under surface pure sharply

defined white. Top of muzzle greyish; sides dark brown, this

colour passing backwards and joining the dark eye-vings. Ears

of medium length, the proectote dull blackish. Hands white, the

dark of the forearms encroaching a little on the metacarpus.

Feet brown over the greater part of the metatarsus; toes and

hallueal border of metatarsus white. Tail very long, fairly well-

haired, not conspicuously tufted, brown for its proximal half

above, then changing gradually to white; underside wholly white,

except just at the base.

Skull in general structure like that of #. excelsior, but larger

throughout ; brain-case rather less convex above.

Dimensions of the type (young adult) :—

Head and body 164 mm. ; tail 248; hind foot 37; ear 26:5.

Skull: greatest length 43; basilar length 35; nasals 16°3; breadth

of brain-case 17°2; palatilar length 19°8; palatal foramina 8:5;

upper molar series 8:2.

The dimensions will at once separate from any of its allies this

fine long-tailed Rat, which I have much pleasure in dedicating to

its discoverer, Mr. Malcolm Anderson.

i MR. OLDFIELD THOMAS ON

30. APODEMUS SPECIOSUS PENINSULEZ Thos.

3. 2202, 2211, 2212, 2213, 2218. 60 miles S.E. of Min-chow,

Kan-su. 8000’,

SG. 2227, 2228, 2231, 2232, 9. 22299" 10 miles S. of Tau=

chow, Kan-su. 8000’—9000’.

6. 2234, 2235, 2237. 9. 2242. Mts. S.E. of Tau-chow,

Kan-su. 10,000’.

3. 2285, 2305, 2308. Wen-hsien Country, 8. Kan-su.

I have already commented on the resemblance of the Shan-si

Field-Mouse to that of Korea, and now findthat those from Kan-

su are again similar in every respect and should undoubtedly be

referred to the same subspecies, the animal therefore presenting

a noteworthy example of the uniformity of the Fauna of China

N. of lat. 34°. As usual, the representative southern form in

Sze-chwan is at once distinguishable by its much darker colour.

31. APODEMUS SPECIOSUS CHEVRIERI M.-Edw.

246,19. 23 miles 8.E. of Ta-tsien-lu, Sze-chwan. 10,000’.

13 6,13 2. Ta-tsien-lu. 9000’.

17 g,11 2. Omi-san. 6000'—-9500’.

. APODEMUS AGRARIUS PALLIDIOR Thos.

2195. Si-Ho-Hsien, S. Kan-su. 5000’.

2292. 2. 2293. Wen-hsien Country, 8S. Kan-su.

SEN

33. APODEMUS FERGUSSONI.

Thos. Abstr. P. Z. 8. 1911, p. 4 (Feb. 14).

Q. 2296. Wen-hsien Country, 8. Kan-su.

@. 2499 (yg.), 2508, 2509 (yg.). Omi-san. 9500’.

Near 4. agrarius, but without the slightest trace of a dorsal

line.

Size and proportions about as in agrarius. Fur short and fine,

without spines in a May specimen, with them in August

examples ; hairs of back about 7 mm. in length. General colour

above uniform drab-brown, not buffy on sides. Under surface

soiled greyish, the bases of the hairs slaty, the tips greyish white ;

lateral line of demarcation fairly well defined. Back without the

slightest trace of a median dorsal line, or even of a median darker

dorsal area, some trace of a line being always perceptible in

A. agrarius. Ears short, coloured like the head. Hands and feet

white. Tail rather shorter than head and body, brown above,

dull whitish below.

Skull about as in 4. agrarvus.

Dimensions of the type :—

Head and body 107 mm.; tail 98; hind foot 24; ear 15.

Skull: greatest length 27-7; breadth of brain-case 12; upper

molar series 4.

ffab. 8. Kan-su and Western Sze-ch wan.

MAMMALS FROM WESTERN CHINA, NY

Type. Adult female, B.M. Now 11.2.1.180. Oviginal number

2296. Collected 8 May, 1910.

This Mouse is readily distinguishable from any of the forms of

agrarius by the entire absence of a dorsal line, some trace of a line

being perceptible in all the available specimens of A. agrarius

ningpoensis, the form in which it is least developed. Hven then

I should like to call it a subspecies of agrariws, but do not

venture to do so without seeing intermediate specimens. The

type was obtained at or near the same locality as a well-marked

example of A. agrarius pallidior (no. 2292),so that in this region

at least there seems to be no intergradation.

I have named this animal after the Rev. W. N. Fergusson,

from whom the Museum has received four specimens of it,

collected at Yin-shin-wan, N.W. of Chong-tu, in 1909.

CRICETULUS ANDERSONI Thos.

3S. 2215, 2216. 9. 2217. 60 milesS.E. of Min-chow, Kan-su.

©. 2225, 2233. 10 miles 8. of Tau-chow, Kan-su.

35. Microtus IRENE.

Thos. Abstr. P. Z.S. 1911, p. 5 (Feb. 14).

. 2383, 2386, 2397, 2400, 2404. 9. 2384, 2385, 2392,

2395, 2396, 2398, 2399, 2401, 2402, 2403. ‘Ta-tsien-lu, W.

Sze- ahem 9000" 12,000’.

Rather larger than WV. arvalis. M, with the same spaces con-

nected as in Pitymys.

Fur soft, fine, rather woolly; hairs of back about 9 mm. in

length. General colour above brown, under surface slaty

washed with greyish white; not sharply defined laterally. Ears

projecting beyond the fur, uniformly brown. A small lateral

gland present on each side in front of the hips in the males.

Hands and feet brownish white ; soles with six pads. ‘Tail about

twice the length of the foot; brown above, paler on sides and

below, not so strongly contrasted as in J. malcolmi. Mamme

2—2 =

Skull of the usual flattened shape, its upper outline straight or

even slightly concave over the orbits. Brain-case remarkably

small in proportion to the face, smooth, rounded, its ridges and

angles scarcely developed ; erale slender, scarcely homed down-

poids interorbital space unusually narrow, smoothly rounded.

iF alatal foramina fairly long. Posterior palate normal, the lateral

pits not specially deep ; mesopter ygoid fossa of normal ior eadth, its

anterior border rounded. Bullee decidedly smaller than in most

species of this size.

Incisors of medium strength, considerably bevelled in front. M?

with pattern very similar to that figured by Blasius in JZ. savii

(Faun. Deuts. fig. 221, p. 394). M, “with only four closed spaces,

the fifth and sixth (counting from behind and including the

posterior transverse space) opening into each other (as in Pity ymys)

174 MR. OLDFIELD THOMAS ON

and also into the seventh, the anterior space, which is curved

inwards to form a fifth internal angle, while there are only three

external angles; the tooth is thus very like that figured by

Bichner in J. limnophilus (Mamm. Przewalsk. pl. xvii. fig. 13).

Dimensions of the type :—

Head and body 98 mm.; tail 37; hind foot 18; ear 13.

Skull: condylo-basal length 24:2; basilar length 22; greatest

breadth 14°8; nasals 7; interorbital breadth 3°8; breadth of

brain-case 12; height of forehead from alveolus of m* 7; palatilar

length 13; palatal foramina 4°8; diagonal diameter of bulla in

‘horizontal plane 6°5 ; upper molar series (crowns) 5-7.

Hab. as above.

Type. Old female, B.M. No. 11.2.1.195. Original number 2396.

Collected 2 July, 1910.

This species is peculiar for the proportionately small size of its

brain-case and its narrow interorbital space. From J/. limnophilus

Biichn., which may possibly be allied to it, itis distinguishable by

having only three internal angles on m*; the shape of the brain-

case would also seem to be more normal in that animal.

36. Microrus MALCOLMI.

Thos. Abstr. P. Z.S. 1911, p. 5 (Feb. 14).

6. 2245, 2250, 2251, 2252. 9. 2246, 2253, 2254, 2255

2956, 2257. Mts. 8.K. of Tau-chow, Kan-su.

A medium-sized Vole with high arched skull, somewhat similar

on a smaller scale to that of MW. calamorwm 'Thos.

Fur soft and fine, of medium length; hairs of back about

9-10 mm. in length *, therefore conspicuously shorter than in

Proedromys. Colour above lined brown, rather lighter than

“bistre”; under surface washed with greyish white, rather

markedly contrasted with the upper colour. Ears slightly

projecting from the fur, brown. Upper surface of hands and feet

dull whitish ; soles with six pads. ‘Tail rather more than double

the length of the foot, brown above, dull white on sides and

below. Mamme apparently 2—2= 8, but this cannot be made out

with certainty.

Skull rather like that of JZ. calamorum on a small scale,

though not so strong and heavily ridged. In general shape it 1s

not flattened above, but high, arched above, the nasal profile

bowed downwards, the waist narrow and proportionately far

forward. Surface everywhere smooth and unridged; squamosal

projections scarcely perceptible. Nasals nearly attaining to the

level of the premaxille behind. JInterorbital space narrow,

smooth, evenly convex above, not flattened or hollowed mesially.

Brain-case smoothly rounded. Palatal foramina of equal breadth

throughout, their length exceeding that of m’+m*. Posterior

palate with very deep lateral pits and unusually narrow meso-

pterygoid space. Bulle decidedly larger than the normal.

* In this measurement I always ignore the outstanding tips of the longer hairs,

and only measure to the ends of the thick mass of under-fur.

MAMMALS FROM WESTERN CHINA. mbes)

Upper incisors narrow, considerably bevelled laterally.

Molars of normal structure, the pattern of the upper ones, so

far as spaces and angles are concerned, essentially as figured by

Blasius (p. 879) in M. arvalis, though the posterior lobe of m?’ is

rather narrower and more elongated. Lower teeth as figured by

the same author (p. 366) in JZ. ratticeps, m, with a very similar

anterior lobe and the same number of closed triangles.

Dimensions of the type :—

Head and body 103 mm. ; tail ay ; hind foot (c.u.) 17; ear 13.

Skull: condylo-basal eat 26°2; basilar length 23°5; greatest

breadth 15; nasals 7; “bale orbital Beeaanh 3° 7; bre: an of brain-

case 12; height of crown from alveolus of m* 9:2; palatilar length

13°25 diastema 8 ie Rene foramina 5; diagonal length of bulla

in horizontal plane 82 ; upper molar series (er owns) 6-1.

Other specimens are smaller, the smallest adult having a

condylo-basal length of 23°4, with a hind foot of 16 mm.

Hab. as above.

Type. Adult male, B.M. No. 11.2.1.199. Original number 2245,

Collected 4 April, 1910.

I can find no described species to which this Vole can be

assigned. Its high skull separates it at once from the great

mass of flat-skulled Voles, the shape more recalling that in

M. calamorum, a species possessing a lateral gland and only five

foot-pads. Satunin’s three species from Zaidam, W. of Kan-su, are

all markedly larger than I. malcolmi. Biichner’s W/. limnophilus

from the same district 1s perhaps more closely allied, but would

seem to have a paler general colour and a much more roughened

and angular skull.

37. Microtus (ANTELIOMYS) CHINENSIS Thos.

6. 2328, 2329, 2332, 2238, 2339. ©. 2326, 2327, 2331,

2333, 2334, 2335, 2340, 2341. 23 miles 8.E. of Ta-tsien-lu.

10,000’.

3. 2512, 2519, 2534, 2543, 2544. ©. 2491, 2492, 2528,

2551, 2553. Omi-san. 9500’.

This striking Vole was described in 1891 * from a specimen

found in the stomach of a snake. No other examples have been

obtained until now.

The type-locality was Kia-ting-fu, quite close to Omi-san.

38. Microtus (CARYOMYS) EVA.

Thos. Abstr. P. Z. 8S. 1911, p. 4 (Feb. 14).

3. 2238, 2239. 9. 2240, 2241. Mts. S.E. of Tau-chow,

Kan-su. 10,000".

[Caryomys,

Like ZHothenomys in external and general cranial characters,

but the teeth with the triangles nearly all closed, instead of being

mostly open and connected with each other. Additional postero-

* Ann, Mag. N. H. (6) viii p. 117, 1891.

176 MR. OLDFIELD THOMAS ON

internal lobes on m' and m* reduced to minute and scarcely

perceptible projections.

Type, Microtus inez 'Thos.

Now that a third species, essentially similar to M. inez and

M. nux, has turned up, I think it advisable to form a special

subgenus for the reception of the three. On first describing

M. inez™ I purposely delayed making a new subgeneric name

until we had evidence as to the existence of other species allied

to that peculiar species, but full details were given as to its

characteristics. |

Allied to WZ. (C.) inez and nux, but with much longer tail.

Form slender, the general proportions, chiefly owing to the

unusually long tail, looking very different from those of the allied

species. Fur long, soft, and fine; hairs of back nearly 10 mm. in

length. General colour above pale brown, greyer than in nux

and iez, the type nearly approaching “ broccoli-brown,” but other

specimens of a warmer tint. Under surface rather variable,

greyish slaty, washed with buffy whitish or drab. Hars about as

long as the fur, and of about the same colour. Hands and feet

white; soles with six pads. Tail remarkably long for a Vole,

nearly as long as the body without the head ; dark brown above,

dull whitish below.

Skull very like that of MZ. nux, except that it is smoother and

more lightly built; but even this may be a question of age.

Teeth almost exactly as in J/. nwx ; second and third spaces of

m*, and first and second of m*, communicating with each other—

all the other triangles closed (except, of course, in the case of m,).

Dimensions of the type :—

Head and body 88 mm.; tail 50; hind foot 16:5; ear 12.

Skull: cody losis length 22°6 ; basilar length 20°5 ; greatest

breadth 13°5; nasals 6°8; interorbital breadth 4; breadth of

brain-case 11: 2: ; paletel length 10°6 ; palatal foramina 4-1; upper

molar series (crowns) 5.

Hab. as above.

Type. Adult male. B.M. No. 11.2.1.223. Original number 2238.

Collected 3 April, 1910.

This Vole is at once distinguishable from all its allies by its

remarkably long tail.

39. Microtus (HorHENOMYS) MELANOGASTER M.-Kdw.

G6. 2284, 2290, 2291; 2297, 2304, 2306, 2307. — Qt 2298,

2299, 2300, 2303. Wen-hsien Country, 8. Kan-su.

3. 2421 (melanoid). 45 miles S.W. of Ya-chow, Sze-chwan.

4000'.

@. 2461. Omi-san. 6000’.

I fail to find any tangible difference between thé Kan-su series

and the two s specimens ‘from Sze-chwan, which latter are nearly

topotypical of the species. ne

* P_Z.S. 1908, p. 976.

MAMMALS FROM WESTERN CHINA. 177

40. PRoEDRomyYS BEDFORDT.

Thos. Abstr. P. Z.8. 1911, p. 4 (Feb. 14).

@. 2214. 60 miles S.H. of Min-Chow, Kan-su. 8000".

11 March, 1910. B.M. No, 11.2.1.235. Type.

PROEDROMYS *.

General external form as in Jicrotus. Fur very long. Sole-

pads 6. Mamme 2—2=8.

Skull heavily built, high, its upper profile strongly curved.

Postorbital squamosal projections strongly developed, almost

peg-like. Palatal foramina long. Posterior palate normal.

Bulle rather large.

Teeth. Upper incisors heavy, strongly curved, not thrown

forward ; their front face grooved on its outer third. Lower

incisors short posteriorly, ending at the bottom of the notch

between the angular and condyloid processes, as in Anteliomys.

Molars with their enamel spaces all completely separated from

each other, the anterior wall of each upper space and posterior of

each lower one unusually strongly curved, so as to form in many

cases nearly half of a circle. Number of spaces and salient angles

as follows :—

M.1 : spaces, 3 inner and 3 outer angles, M. 2/ 4-2: 3.M. 3 4.2. ane

5 4 5.3 3.3.

M? very peculiar, consisting of the usual transverse, second and

third spaces, and then acircular posterior lobe directed externally,

the posterior lobe of this tooth in every other Vole, so far as I am

aware, being directed inwards. Anterior space of m, forming

a simple crescent. M, with broad connected spaces across it as

usual.

I fail to find any group into which this lonesiemed Vole can be

fitted. Its heavy bowed skull, grooved incisors, peculiar curved

walled tooth-spaces, and curious m, seem together to distinguish

it from any described genus. The grooving of the incisors is

possibly an individual character, but the other features of the

animal are so marked that it should certainly have a special

name.

PROEDROMYS BEDFORDI.

Size medium. General form that of an average Vole, the tail

rather more than a third the length of the head and body. Fur

excessively long, soft, and fine; hairs of back about 16 mm. in

length. General colour above coarsely lined dull brown (darker

than ‘ broccoli-orown”), the lower flanks more drabby ; under

surface slaty drab washed with brownish white. Ears hairy, pale

* apdetpos, a President. Named in honour of the Society’s President, to whose

generosity the discovery of this animal is due.

Proc. Zoon, > oc.—1911, No, XII, 12

178° WR. OLDFIELD THOMAS ON

brown. Hands and feet dull white. Tail well-haired, brown

above, dull white on sides and below.

Skull and teeth as indicated above.

Dimensions: of the type :—

Head and body 103 mm.; tail 41; hind foot 18; ear 13.

Skull: basal length (c.) 26; greatest breadth 16; nasals

7-6 x 3:2; interorbital breadth 3-6; breadth of brain-case 12°2 ;

palatilar length 13°7; diastema 8; palatal foramina 6; upper

molar series (crowns), 6°8..

Hab. and Type as above.

Interesting and peculian as this species is in essential characters,

i6 is externally a very ordinary-looking Vole, and indeed scarcely

distinguishable from Microtus matcolmi, which Mr. Anderson found

im some numbers in the same region..

41,. Myospauax cAnsus Lyon.

3. 2263, 2272, 2274. ©&.. 2262, 2264, 2265, 2266, 2267,

9968, 2271, 2275, 2276.. 10 miles 8. of Tau-chow, Kan-su. 8500".

These are practically topotypes of Dr. Lyon's species, and a

study of them shows that the series obtained in Shen-si by

Mr. Anderson, and previously referred to J/. cansus*, should

be subspecifically separated.

The form may be called :—

MyesPALAX CANSUS: SHENSELUS..

Thos.. Abstr. P..Z.S. 1911, p..5 (Feb. 14).

General characters as in true cansus, but the colour warmer and

the tooth-row longer..

Taking only full-coloured specimens into consideration the

coloured ends of the hairs above approach “ ochraceous-buff,”

while in cansus they average distinctly paler, approximating to

“pinkish-buff.” As to the tooth-row, among seven adult specimens

of true cansus (including three well-grown males) the alveolar length

of the upper series never exceeds 10-0 mm., while of 16 specimens

of shenseiws some (males) may almost reach 12-0 mm., while nearly

all, including females, are 11:0 and upwards. One fully adult

specimen only, a female, has the dimension 10:0.

Dr. Lyon’s type, a male (although originally called a female),

seems to have been unusually large, as its tooth-row measures

about that of a female shenseiws, and therefore more than any

of our specimens of the Kan-su form. This would, however,

appear to have been an exceptional specimen. i

Dimensions of the type :—

Head and body 178 mm.; tail 54; hind foot 30.

Skull: condylo-basal length 45; zygomatic breadth 34; nasals

18x7:7; interorbital breadth 6:5; palatilar length 23:5; upper

molar series (crowns) 11:4, (alveoli) 11:9.

Hab. Shen-si. Type from Yu-lin-fu.

* PZ. 8. 1908, p. 978.

MAMMALS FROM WESTERN CHINA- 179

Type. Old male. B.M. No. 9.1.1.216. Original number 1800.

Collected 30 April, 1908.

Dr. Allen’s “ Myotalpa” rufescens* would seem also to have

the small teeth of true JZ. causus, and is geographically closer to

the latter.

42. Rarzomys vestizus M.-Edi,

©, 2559. Omi-san. 9500’.

The relationship of this form to Gray’s #. sinensis’ has not

hitherto been satisfactorily defined. It seems, however, to be

readily characterized by the much larger size of the skull, the

great development of the cranial crests, the length and softness of

the fur, and the shortness of the tail. . sinensis probably came

from somewhere in South China, perhaps the neighbourhood of

Canton.

A third Chinese species: may be described as follows :—

LHIZOMYS DAVIDI.

Thos. Abstr. P. Z.S. 1911, p. 5 (Feb. 14).

Colour and character of fur, and length of tail, about as in

RF. vestitus ; but size much smaller, nearly as in A. sinensis.

Skull shghtly larger than that of 2. sinensis, the male 2. sinensis

about equaling the female &. davidi. Crests rather more

strongly developed, comparing like sex with like. Nasals more

narrowed behind and premaxillary processes proportionately

broader, so that, while in &, sinensis the combined breadth behind

of the two nasals about equals either fronto-premaxillary suture;

in 2. davidi the length of each suture is about twice the distance

that separates their nearest points: in the middle line. Bulle

larger.

Dimensions of the type, measured in- skin :—

Head and body (c.) 350 mm.; tail 69; hind foot (wet) 42; ear

(wet) 15.

Skull: condylo-basal length 66.; upper length from lambda to

tip of nasals 53-5; greatest breadth 49°5; nasals 23°2 x 8:4:;.

fronto-premaxillary suture 6°5 ; interorbital breadth 9°2; greatest

occipital breadth 30; palatilar length 35; diastema 20;. upper

molar series (crowns) 13:3.

An old male skull measures 49 mm..in upper length.

Hab. Kuatun, N.W..Fokien. 3500’.

Type. Old female. B.M. No: 96.12.1.6.. Original number 9.

Collected December 1898, and presented by J. de La Touche, Esy.-

Eleven specimens examined.

This fine species, of which the Museum. possesses:a good series,

presented by Messrs. F. W. Styan, J. de La Touche, and C. B.

Rickett, is distinguishable from 7. sinensis by its soft fur, shorter

tail, and the above-mentioned cranial characters. From &. restitus,

again, by its very much smaller size.

* Bull. Am. Mus. N. H. xxvi. p. 429, 1909.

180 ON MAMMALS FROM WESTERN CHINA.

It is with great pleasure that I name a Chinese animal after

Pere David, the famous French naturalist and collector, to whom

Science owes the discovery of the wonderful series of animals

on which Milne-Edwards’s ‘ Recherches Mammiféres’ was based.

Kuatun, the Jocality of Rhizomys davidi, was itself discovered as

a collecting-station by Pere David, though we owe its more recent

exploitation to our own countrymen.

43. LEPUS SECHUENENSIS de Wint.

6. 2258, 2261. ©. 2259. 30 miles N.W. of Tau-chow,

Kan-su. 11,000’.

Although described as from Sze-chwan, it is probable that this

species really belongs to the fauna N. and N.W. of the T'sin-ling

range. The type was said to be from Worth-western Sze-chwan,

and that part of the Province extends into the faunistic region

of which Kan-su is a part. No Hare is as yet known from the

typical forested area of Western Sze-chwan.

44, OcHorona cANsA Lyon.

SO. 2236, 2243, 2248, 2249. ©. 2226, 2244, 2247. Mts.S.E.

of Tau-chow, Kan-su.

Tau-chow (‘Tao-cheo) is the type locality of Dr. Lyon’s species.

45. OcHOTONA TIBETANA, M.-Kdw.

3. 2325. 9. 2324, 23 milesS.E. of Ta-tsien-lu, Sze-chwan.

10,000’.

eovaiaie UWrasemelin, GOO,

3. 2525. Omi-san. 9500’.

These specimens indicate that O. tabetana has a wide range of

colour-variation, very much as in the allied O. hodgsont.

46. CAPREOLUS BEDFORDI Thos.

3S. 2208, 2205, 2207, 2210. 9. 2204, 2208, 2209. 60 miles

S.HE. of Min-chow, Kan-su. 8000’.

47. Moscuus stranicus Bichn.

2 (immature). 2223. 10 milesS.of Tau-chow, Kan-su. 8500’.

The first example of this striking species that has come to the

British Museum.

48. GAZELLA sp.

3 (young). 2260. 30 miles N.W. of Tau-chow, Kan-su. 11,000’.

No. 87.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

November 15th, 1910.

Dr. 8. F. Harmer, M.A., F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Srecrerary read a Report on the additions that had been

made to the Society's Menagerie during the months of June,

July, August, September, and October 1910.

Mr, J. Lewis Bonuote, M.A., F.L.S., F.Z.S., exhibited a spirit-

specimen of a young Cairo Spiny Mouse (Acomys cahirinus),

about twelve hours old, pointing out the advanced state of the

young at birth as compared with the common House-Mouse, and

remarked that the period of gestation in Acomys was eleven days

at the most, and that the usual number in a litter was three.

Mr. Bonnore also remarked on a pair of hybrids between the

Bramble-Finch (“ringilla montifringilla) and the Chaffinch (frin-

gilla celebs), which he had deposited at the Gardens and which

represented a cross bred for the first time in 1907. He drew

attention te the resemblances in the plumage of these birds, and

stated that the fertility of the hybrids had not yet been tested.

Mrs. R. Hata Tuomas, F.Z.S., exhibited a series of skins

illustrating an experiment in Pheasant-breeding, and gave an

account of the production of Thawmalea obscura in the F,

generation from a cross between 7. amhersti 9 and TZ. picta S.

Dr. W. Nicotu and Prof, E. A. Mincutn, M.A., V.P.Z.S., exhi-

bited specimens of two species of Cysticercoids found in the body-

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Stxpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in adyva ice.

cavity of rat-fleas (Ceratophyllus fasciatus). One of these species

had been found in about four per cent. of the fleas examined and

had been shown, by means of feeding experiments carried on by

Dr. Nicoll, to be the larval form of the common rat-tapeworm

(Hymenolepis diminuta). The other species, of which so far only a

single example had been found, was probably the larva of Hyme-

nolepis murind, a species occurring in rats and mice, and very

similar to, possibly identical with, H. nana, a dangerous tapeworm

of man. The chief interest of the discovery lay in the fact that

no intermediate host was previously known for this tapeworm.

The Hon. N. Cuarites Roruscuitp, M.A., F.Z.S., exhibited

several species of Fleas which were of interest in connection with

the spreading of plague by these insects.

Mr. R. I. Pococr, F.L.S., F.Z.8., exhibited a living specimen

of the Black Rat (J/us ratius) from Sark, which had recently

been presented to the Society’s Menagerie by Mrs. C. Russell.

Mr. R. I. Pocock also exhibited a female hybrid, bred in the

Society’s Gardens, between a male Black Lemur (Lemur macaco)

and a female of the Red-fronted variety of the Fulvous Lemur

(Lemur fuluus rufifrons), and pointed out that the offspring re-

sembled neither of its parents. It had not inherited the facial

fringe of its father nor the white over the eyes and on the fore-

head of its mother, this area of the head being dark ashy black

with a deeper tinted central line, more approaching the colour

seen in the Black-fronted variety of the Fulvous Lemur (ZL. fulvus

nigrifrons).

Mr. J. Lewis Bonnore, M.A., F.LS., F.Z.S., read a paper

dealing with some experiments he had made on the occurrence of

the webfoot character in Pigeons. After referring to Mr. R.

Staples Browne’s paper on the subject in the P. Z.8. for 1905, in

which that gentleman had shown the webfoot to be a. simple

Mendelian recessive, Mr. Bonhote instanced further cases from

the lofts of Mr. F. W. Smalley, F.Z.S., that bore out Mr. Staples

Browne’s conclusions. Both these gentlemen, however, had been

kind enough to give the author birds from their strains, and in

the first instance when webbed birds from the different strains

were crossed an irregular result—namely, 4 normal and 1 webbed

—was obtained. Matings from these birds were continued, and

the results were, in almost every case, contrary to Mendelian

expectations, normals throwing webs, and webs throwing normals.

After discussing various suggestions, Mr. Bonhote came to the

conclusion that no really satisfactory explanation was forthcoming.

The Mendelian inheritance was apparently there, but dominated

and modified by some other agency, and he had been able to find

no single explanation which would cover all the results.

Mr. Epwarp Decen, F.Z.8., read a paper entitled “ Notes on

the little-known Lizard Lacerta jacksoni Blgr., with special re-

ference to its Cranial Characters,” based on a series of specimens

recently procured by Mr. R. Kemp in British East Africa.’

Mr. G. A. Boutenerr, F.R.S., V.P.Z.S., read a paper ‘“ On

Lacerta peloponnesiaca Bibr.,” which contained a new description

of this little-known lizard, made from living specimens in the

Society's Gardens, with a view to fixing its correct position in the

genus Lacerta.

A paper entitled “‘ Remarks on two Species of Fishes of the

genus Gobius, from observations made at Roscoff,” was read by

Mr. HK. G. Boulenger, communicated by Mr. G. A. Boulenger,

F.R.S., V.P.Z.S., dealing with the specific distinction of Globius

minutus and %. microps.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, November 29th, 1910, at half-past Eight

o'clock p.m., when the following communications will be

made :—

1. Dr. H. B. Fantuam, B.A., F.Z.8., and H. Hammonp Smrrx,

M.R.CS., L.R.CP ZS.

On a Possible Cause of Pneumo-enteritis in the Red Grouse

(Lagopus scoticus).

2. F. E. Bepparp, M.A., F.BS., F.Z.S.

On the Alimentary Tract of certain Birds, and on the

Mesenteric Relations of the Intestinal Loops.

3. Prof. A. CABRERA, C.M.Z.S.

On the Specimens of Spotted Hyzenas in the British Museum

(Natural History).

Pee Died: Ei Gemuitt, M.A., D.Se.

The Development of Solaster endeoa Forbes.

SS ee ee |

The following communications have been received :—.

1. Grorce P. FARRAN.

Plankton from Christmas Island, Indian Ocean.—I. On

Copepoda of the Family Coryczide.

2. W. N. F. Woopianp, F.Z.8.

On the Structure and Function of the Gas-Glands and Retia

Mirabilia associated with the Gas-Bladder of some Teleostean

Fishes, with notes on the Teleost Pancreas.

3. H. R. Hoge, M.A., F.Z.S.

On some New Zealand Spiders.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZooLoGiIcaL Society oF Lonpon,

RecEnt’s Park, Lonpon, N.W.

November 22nd, 1910.

MOTOR ‘BUSES FOR THE SCIENTIFIC MEETINGS.

On the night of each Scientific Meeting Motor ’Buses

@ will run direct to the Gardens from Regent’s Park Tube

= Station, starting at 8.5 and 8.20 p.m. The return

A buses after the Meeting will leave the Gardens at 10.15

gand 10.35 p.m. These ’buses will be labelled

|PRIVATE and no fare will be charged to Fellows

EH attending the Meetings.

No. 88.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

November 29th, 1910.

Dr. Henry Woopwarp, F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. D. Sera-Smiru, F.Z.8., Curator of Birds, exhibited, through

the kindness of Mons. Pauvvels, a blue variety of the Budgerigar

(Melopsitiacus undulatus), as well as a yellow variety and a

normal green specimen.

The two varieties showed distinctly the splitting up of the

component coloration, the blue bird lacking all trace of yellow

pigment, while in the yellow variety the blue colouring of the

normal bird had disappeared.

Dr. Witt1Am Nicott, of the Lister Institute of Preventive

Medicine, gave a demonstration of his method for the collection

of Trematodes.

Dr. R. T. Lereer, F.Z.8., exhibited two photographs and

some specimens showing Onchocerciasis in Beef imported from

Queensland.

Dr. H. B. Fanruam, F.Z.8., and Dr. H. Hammonp Smrrtu,

F.Z.S., contributed a paper ‘ On a Possible Cause of Pneumo-

enteritis in the Red Grouse (Lagopus scoticus).” The authors

recorded that in Grouse-chicks dying of Coccidiosis, many of

which showed symptoms of pneumonia, they found Coccidian

odcysts in the bronchioles, bronchi, and trachea. The Coccidian

cysts in the bronchioles were probably capable of setting up

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance.

sufficient irritation to account for the pneumonic symptoms.

These observations were interesting as showing that the much

criticised views of Klein, Tegetmeier, and others on “ pneumo-

enteritis” as a cause of mortality in Grouse may have some

foundation in fact.

Dr. J. F. Gemiin, M.A., D.Sc., Lecturer on Embryology in the

University of Glasgow, gave an account, illustrated by lantern-

slides and specimens, of his memoir on “The Development of

Solaster endeca Forbes,” communicated to the Society by Prof.

J. Arthur Thomson, F.Z.8.

The author described the ovaries and ova and the processes

of spawning, fertilization, segmentation, and gastrulation, and then

dealt with the characters of the free-swimming larve and the

changes related to the metamorphosis. He discussed the develop-

ment of the internal cavities and of the skeleton, and described

the methods he had employed in obtaining and rearing the larve.

The memoir, in addition to details of adult anatomy, contained

a description of the following points in development :—

Segmentation total equal; blastula by egression ; gastrula by

invagination ; early closure of blastopore ; no larval mouth.

Free-swimming larva with three arms and a muscular sucker ;

attachment by sucker.

Metamorphosis such that while in point of external form the

left side of the larva becomes the oral surface of the starfish, in

reality the epiderm of the oral surface is derived in great part from

that of the anterior part of the early bilateral larva, and conversely

the aboral epiderm is derived chiefly from that of the posterior end

of the larva.

Archenteron dividing into anterior and posterior vesicles and

middle chamber or enteron. Anterior vesicle giving rise to pre-

oral, axial, and epigastric cceeloms, hydroccele, dorsal sac, internal

oral sinus, and part of external oral sinus. Posterior vesicle

giving rise to hypogastric and pharyngeal cceloms, rest of external

oral sinus, aboral circular sinus, and genital rachis.

The full set of hydroccele pouches completed slowly, the series

running in the watch-hand direction as seen from the oral side ;

opening of stone-canal between radii I and IL; anus in inter-

radius V/VI.

Terminal plates double; rest of aboral plates showing atypical

distribution.

A larval nervous system and a statolith-like body in the

posterior ccelom.

Mr. F. BE. BeppArp, M.A., F.R.S., F.Z.8., Prosector to the

Society, presented a paper ‘“‘On the Alimentary Tract of certain

Birds, and on the Mesenteric Relations of the Intestinal Loops,”

based on notes he had accumulated relative to the viscera of

Birds which had died in the Society’s Gardens. The paper dealt

more particularly with species that had not been carefully studied

from the point of view of the convolutions of the intestine, and

attention was called to a considerable series of Birds.

Mr. OtprievD THomas, F.R.S., F.Z.S., communicated a paper

by Prof. Ancet Caprera, C.M.Z.8., “On the Specimens of Spotted

Hyenas in the British Museum (Nat. Hist.),” in which he de-

scribed three apparently new forms.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, December 13th, 1910, at half-past Hight

o'clock P.mM., when the following communications will be

made :—

1. ZootocicaAL NOMENCLATURE.

Dr. W. K. Hoyle, M.A., F.Z.8., will explain the Report of

the International Commission on Zoological Nomenclature pre-

sented to the Graz Meeting of the International Zoological

Congress, 1910, and will refer in particular to the proposals

made for the protection of well-known zoological names.

. E. 8. Goopricy, M.A., F.R.S., F.Z.S.

On the Segmentation of the Occipital Region of the Head in

the Batrachia Urodela.

3. Dr. W. N. F. Woopuanp, F.Z.S.

On the Structure and Function of the Gas-Glands and Retia

Mirabilia associated with the Gas-Bladder of some Teleostean

Fishes, with notes on the Teleost Pancreas.

4. OLDFIELD THomas, F.R.S., F.Z.8.

The Mammals of the Tenth Edition of Linneus: an Attempt

to fix the Types of the Genera and the exact Bases and Localities

of the Species.

The following communications have been received :—

1. Grorcr P. Farran.

Plankton from Christmas Island, Indian Ocean.—I. On

Copepoda, of the Family Coryceidee.

2. H. R. Hoae, M.A., F.Z.S8.

On some New Zealand Spiders.

3. Dr. F. D. Wetca, F.Z.8.

Observations on different Species of Hylobates now or recently

living in the Society’s Gardens, and a Symphalangus syndactylus,

with Notes on Skins in the Natural History Museum.

4. R. LyDEKKER.

On Tragelaphus buatoni.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL SocrETY OF LONDON,

Recent’s Park, Lonpon, N.W.

December 6th, 1910.

MOTOR 'BUSES FOR THE SCIENTIFIC MEETINGS.

1 On the night of each Scientific Meeting Motor ?Buses

| will run direct to the Gardens from Regent’s Park Tube

, Station, starting at 6.5 and 8.20 p.m. The return

| ‘buses after the Meeting will leave the Gardens at 10.15

Hand 10.35 p.m, These *buses will be labelled

Ww PRIVATE and no fare will be charged to Fellows

Y attending the Meetings.

No. 89.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

December 13th, 1910.

G. A. Boutencer, Esq., F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary read a Report on the additions that had been

made to the Society’s Menagerie during the month of November

1910.

Dr. H. Hammonp Surru, F.Z.S., exhibited a mounted specimen

of a male Red Grouse, from Abington, Lanarkshire, which dis-

played a curious variety of the ordinary plumage of this species.

Mr. D. Sers-Smiru, F.Z.S., Curator of Birds, exhibited some

skins of the Australian Yellow-rumped Finch (Munia flavi-

prymna). ‘These birds had been kept alive in an outdoor aviary

in England, and had developed certain markings tending towards

those of another closely allied species, Muna custaneithorax.

The exhibitor attributed this to the fact that the former species

was a desert form of the latter, and when placed in a humid

environment tended to revert to the plumage of the latter. He

referred to a paper he had published on this subject in the

‘ Avicultural Magazine,’ 1907, p. 195.

Mr. Epwin S. Goopricu, M.A., F.R.S., F.Z.S., read a paper

“On the Segmentation of the Occipital Region of the Head in

the Batrachia Urodela,” based on his studies of the development

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shil/ings per annum, payable in advance.

of the head region of the Axolotl (Amblystoma tigrinwm). The

head of the Axolotl contained. three segments behind the auditory

capsule. Three metaotic somites were developed in these

segments, of which the first soon disappeared, and the second and

third contributed to the formation of the temporal musele. To

the first segment belonged the glossopharyngeal nerve; to the

next two the vagus. The occipital condyles were developed

between the third and fourth somites. The two hypoglossal

voots corresponded to the fourth and fifth metaotic segments, and

passed out in front of the first and second vertebre. In the

Amniota all these segments were included in the head. ‘The

skull of an Amphibian was thus shorter than that of a Mammal,

yet the condyles were homologous in the two animals. ‘The

shifting backwards or forwards of the condyles was brought

about, not by the inter- or excalation of segments, but by a

tr ansposition from one segment to another. ‘The shifting of the

condyles was comparable to the tr ansposition of the limbs on the

trunk-segments.

Mr. Ouprirtp Tomas, F.R.S., F.Z.8., read a paper entitled

“The Mammals of the Tenth Edition of Linneus: an Attempt

to fix the Types of the Genera and the exact Bases and Localities

of the Species.” It was shown that by the use of tautonymy the

types of nearly all the Linnean genera could be definitely fixed,

the conclusions arrived at by this means agreeing in most cases

with common usage. The type of Simia, however, would not be

S. satyrus but S. syli vand, and of Dasypus D. novemcinctus instead

of D. seacinctus, the consequences of which changes were pointed

out.

Pygathrix, as represented by the two species memeuws and

nigripes, was Shown to be generically distinct from Preshytis, so

that the latter name still remained available for the ordinary

Langurs.

Changes in specific names, due to a complete examination,

were shown to be less numerous than might have been expected,

while the stability of mammalan nomenclature was much

increased by avoiding the danger of what such an examination

might lead to.

Type localities, derived from the original authors quoted by

Linneus, were defined for a considerable number of the species.

Dr. W. E. Hoyts, M.A., F.Z.8., English Member of the Inter-

national Commission on Zoological Nomenclature, explained the

following Report presented to the Graz Meeting of the Inter-

national Zoological Congress, and referred in particular to the

proposals made for the protection of well-known zoological

names.

A discussion followed on the portion relating to the formation

of an Official Listof most frequently used Zoological Names. The

feeling of the Meeting was very strongly in favour of the Inter-

national Congress giving its authority to the formation of a List.of

Zoological Names, the significance of which should not be. altered

by application of the miles of the International Code. It was

unanimously agreed to accept the action of the Congress if it

would adopt this course.

REporT oF THE INTERNATIONAL COMMISSION ON

ZooLegicAL NOMENCLATURE.

FINANCIAL AID FROM THE SMITHSONIAN INSTITUTION.— Owing to

the amount of clerical work connected with the studies conducted

by the Commission, it has been found very difficult in the past

for the Commission to render its decisions as promptly as desirable.

This difficulty has now been overcome by the generous grant of

the sum of $2700 by the Smithsonian Institution; said sum is

available at the discretion of the Commission at any time during

the three years following the grant.

In addition, the Smithsonian Institution has placed at the

disposal of the Commission the sum of $500 to be used in pub-

lishing the “Opinions” rendered by the Commission in its

function as a Court of Appeal. An arrangement has been made

between the Secretary of the Smithsonian Institution and the

Secretary of the Commission, whereby the ‘ Opinions” will be

published by the Institution and forwarded to 1100 libraries, to

the Members of the International Zoological Congress, and to a

limited list of specialists.

OPINIONS RENDERED.—Since October, 1909, the Commission

has rendered 23 “ Opinions” (Nos. 6-28), which are now in press

and which will soon be sent to all Members of the Congress. A

number of cases are still before the Commission for study and

will be passed upon in the near future. The summaries of

Opinions 6-28 are as follows :— ;

6. In case of a genus A, Linneus, 1758, with two species Ab and Ac.—When

a later author divides the genus 4, species 4) and dc, leaving genus A

only species 46, and genus C, monotypic, with species Ce :

The second author is to be construed as LETH fixed the type of the

genus 4, [See Article 30.]

Vote: Affirmative 14; negative 0; not voting 1.

7. On the interpretation of the expression “n. g.,n. sp.” under Article 80 (a).—

The expression ‘“n. g., n. sp.,” used in publication of a new genus for

which no other species is otherwise designated as genotype, is to be

accepted as designation under Article 30 (Oy

Vote: Affirmative 8; negative 3; not voting 2; vote both ways 2.

8. On the retention of ti or iin specific patronymic names, under Article 14 (c)

and Article 19.—Specifie patronymics originally published as ending in

ii (as schrankii, ebbesbornit) are, according to Article 19, to be retained

in their original form, despite the provision of Article 14 (c), which

provides that they should have been formed with only one @,

Vote: Affirmative 11; negative 1; not voting 2; vote both ways 1.

9. The use of the name of a composite genus for a component part requiring

a name.—The decision as to whether the name of a composite genus,

when made up wholly of older genera, is tenable for a component part

requiring a name, depends upon a variety of circumstances. There are

circumstances under which such name may be used, others under which

it may not be used. (Article 30.)

Vote: Affirmative 13; negative 0; not voting 2.

10. Designation of genotypes for genera published with identical limits—If

two genera with the same limits are formed independently by different

authors, without designation of genotypes, any subsequent author may

designate the genotypes (Art. 80g); and if the types designated are not

specifically identical, the two generic names may (other things being

equal) be used for restricted genera containing the types in question.

(Article 25.)

Vote: Affirmative 9; negative 4; not voting 2.

Il. The designation of genotypes by Latreiile, 1810.—The “ Table des genres

avec Vindication de Vespéce qui leur sert de type,” in Latreille’s (1810)

‘ Considérations générales,’ should be accepted as designation of types

of the genera in question. (Article 30.)

Vote: Affirmative 12; negative 1; not voting 2.

12. Stephanoceros fimbriatus (Goldfuss, 1820) vs. Stephanoceros cichhornis

Ehrenberg, 1832.—The generic name Stephanoceros, 1852, is to be used

in preference to Coronella, 1820 (pre-occupied, 1768); the specific name

jfimbriatus, 1820, takes precedence over eichhornii, 1832, which is ad-

mittedly (Khrenberg, 1832 b, 125, and 1838 a, 400-401) jimbriatas, 1820,

renamed. Ehrenberg was right in rejecting Corenel/a, 1820, but in

error in rejecting jumbriatus, 1820; no reason is apparent for per-

petuating his error.

Vote: Affirmative 14; negative 0; not voting 1.

13. The specific name of the Sand-Crab.—Catesby’s (1743) pre-Linnean name

arenarius is not available under the Code, although “reprinted” in

1771; quadratus, 1793, is stated to be pre-occupied; albicans, 1802,

being the next specific name in the list becomes valid, under the

premises submitted.

Vote: Affirmative 12; negative 0; not voting 3.

14. The type species of Ktheostoma Rafinesque, 1819.—The designation of

Li. blennioides Rafinesque, 1819, as type of Htheostoma Rafinesque, 1819,

by Agassiz, 1854, is not invalidated by the fact that Agassiz used as basis

for his generic diagnosis characters taken from an erroneous specific

determination of 1839. Not only does Agassiz distinctly state that

“ Hth. blenniotdes Raf.” is type of ‘“ Htheostoma Raf.,” but even if the

question of the erroneous identification of H. blennioides by Kirtland be

taken into consideration, the conclusion must be drawn that this erroneous

identification did not exclude the original specimens of E. blennioides

from being covered by this specific name; on the contrary, the name

as used by Kirtland, 1839, still involved the type specimens; removing

now the erroneously determined specimens of 1839, which by article

30.e (a) are excluded from consideration in designating the genotype,

the original type specimens of 1819 remain and, upon the premises

submitted, represent the type of the genus.

Vote: Affirmative 9; negative 4; not voting 2.

15. Craspedacusta sowerbii Lankester, 1880, n. g., n. sp., vs. Limnocodium

victoria Allman, 1880, n.g., n. sp., a@ freshwater Medusa.—Craspeda-

custa sowerbit Lankester, 1880, June 17, has clear priority over Limno-

codium victoria Allman, 1880, June 24. Presentation of a paper before

a scientific society does not constitute publication in the sense of the

Code. The Commission is without authority to sanction usage in con-

travention of the provisions of the Code.

Vote: Affirmative 15; negative 0.

16. The status of prebinominal specific names (published prior to 1758) under

Art. 30 d.—In deciding whether a case of absolute tautonymy is present

(under Art. 30d), the citation of a clear prebinominal specific name in

synonymy is to be construed as complying with the demands of

Art. 30d. Examples: Equus caballus (Equus cited in synonymy in the

sense of “the horse”), Alca torda (Alca cited in synonymy in the

sense of ‘the Alca”).

Vote: Affirmative 10; negative 2; not voting 3.

17. Shall the genera of Weber, 1795, be accepted ?—Weber’s ‘Nomenclator

Entomologicus,’ 1795, complies with the requirements of Article 25;

hence the genera in question are to be accepted, in so far as they

individually comply with the conditions of the Code.

Vote: Affirmative 12; negative 1; not voting 2.

18. The type of Hydrus Schneider, 1799.—On basis of the premises, caspzus

Schneider, syn. hydrus Pallas, is type of Hydrus Schneider, Art. 30 d.

Vote: Affirmative 14; negative 0; not voting 1.

19, Plesiops vs. Pharopteryx.—From the evidence, it is not clear that this

case is one of nomenclatorial rather than zoological nature. So far as

the evidence goes, the question as to whether Riippell was in error in

accepting Plesiops as identical with Pharopteryx must be answered from

a systematic point of view. If from our present-day conception of

generic limits, Riippell was correct, no reason is apparent for not

accepting his nomenclatorial decision.

Vote: Affirmative 11; negative 1; not voting 3.

20. Shall the genera of Gronow, 1768, be accepted ?—Gronow, 1763, is binary,

though not consistently binominal. Article 24 demands that an author

be binary, and Article 2 demands that generic names be uninominal.

Under these Articles, Gronow’s genera are to be accepted as complying

with the conditions prescribed by the Code to render a name available

under the Code.

Vote: Affirmative 11; negative 1 ; not voting 3.

21. Shall the genera of Klein, 1744, reprinted by Walbauwm, 1792, be

accepted ?—When Walbaum, 17$)2, reprinted in condensed form (but

did not accept) the genera of Klein, 1744, he did not thereby give to

Klein’s genera any nomenclatorial status, and Klein’s genera do not

therefore gain availability under the present Code by reason of being

quoted by Walbaum.

Vote: Affirmative 12; negative 0; not voting 3.

22. Ceraticthys vs. Cliola.—Whatever Baird’s original intentions may have

been, he and Girard originally published (1853) Ceraticthys as a mono-

typic genus, describing the genotype (C. vigilar) and giving no indica-

tion that there were any intentions other than to publish a “n.¢g.,

”

n.sp.” Under Article 30c¢, vigilax is the type of Ceraticthys.

Vote: Affirmative 12; negative 0; not voting 3.

23. Aspro vs. Cheilodipterus, or Ambassis—Under the premises given,

Centropomus macrodon may be taken as type of Aspro, 1802, and this

generic name suppressed assynonym of Cheilodipterus, thus safeguard -

ing Ambassis.

Vote: Affirmative 8; negative 1; not voting 6.

24. Antennarius Commerson, 1791, and Cuvier, 1817, vs. Histrio Fischer,

1813.—Antennariuvs Commerson is an uninominal generic name (Art. 2)

of an author who used a binary (Art. 25) (though not binominal)

nomenclature. It received nomenclatorial status by virtue of its

publication by Lacépéede, 1798, and should date from that time instead

of frum Cuvier, 1817. It is therefore not necessary to suppress it im

favour of Histrio, 18138.

Vote: Affirmative 12; negative 0; not voting 3.

25. Damesiella Tornquist, 1899, vs. Damesella Walcott, 1905.—Under

Article 36, Recommendations, it is not necessary to reject Damesella,

1905, because of the existence of Damesiel/a, 1893 (1899 ?).

Vote: Affirmative 11; negative 1 ; not voting 3.

26. Cypsilurus vs. Cypselurus.—In view of the number of typographical

errors in Swainson, 1838 aud 1839, the Commission is of the opinion

that Cypsilurus is an evident typographical error and should be corrected

to Cypselurus.

Vote: Affirmative 10; negative 1; not voting 4.

27. Ruppelia and Rupellia vs. Riippellia.—Since a typographical error is

evident, Ruppelia and Rupellia should be corrected to Lvuippellia.

Vote: Affirmative 9; uegative 1; not voting 9.

28. Shall the ‘ Nouvelle Classification’ of Meigen, 1800, be given precedence over

Meigen’s ‘Versuch, 1803 ?—The generic names contained in Meigen’s

‘ Nouvelle Classification,’ 1800, must take precedence over those in his

‘Versuch, 1803, in every case where the former are found valid under

the International Code.

Vote: Affirmative 11; negative 0; not voting 4.

OrrictaAL List OF MOST FREQUENTLY USED ZoontocicaL NAMES.—

There is a desire on the part of some zoologists that certain

very commonly used zoological names should be excepted from

the application of the Law “of Priority, and a proposition to this

effect has been presented to the Commission from the British

Association for the Advancement of Science and the Eastern

Branch of the American Society of Zoologists. That this desire

is so widespread and so deeply rooted as is assumed by some of

our colleagues has not been confirmed by inquiries made by

several members of the Commission. Further, an effort made by

the Secretary to collect from zoologists the most commonly used

and most important generic names has as yet met with such poor

success, that the conclusion does not seem entirely unjustified

that some of our colleagues who may be in favour of such a list

are not as yet sufficiently enthusiastic over the proposition to

induce them to demonstrate their desire by placing into the

hands of the Commission the data upon which such a list must

of necessity be based. Further, there are many colleagues who

are known to us to be directly and enthusiastically opposed to

such list.

After careful consideration of the subject and of the many

difficulties involved, the Commission has decided to propose

to the Congress the trial of a proposition which it is hoped

{

will meet with the approval of both sides of the controversy,

namely :—

(1) The Commission invites all zoslogists to send to the Se eee of the

Comission, prior to November 1, "1910, a list of 100 zoological generic

names which they consider should be studied in connection with the

preparation of an ‘‘ Official List.” Each name should be accompanied

either by the name of the author of the generic name, or by an indica-

tion of the group to which it belongs,

(2) All systematists are invited to send a separate list of the 50 to 100

generic names in their specialty which they look upon as the most im-

portant and most generally used. Hach name should be : oe anied

by the full and complete original bibliographic reference, by the name

of the type species, deter mined according to Art. 30 of the Inter-

national Rules, and by the name of the order and family to which the

genus belongs

(8) All zoologists and paleontologists who give courses in General Zoology

are inyited to supply the Secretary with a list of the text-books

used in said courses, so that said books may be indexed for generic

names.

(4) The Commission will alphabetize all the generic names sent in and will

endeavour, according to circumstances, to determine which are the 100

to 500 most commonly quoted genera.

(5) The genera selected will be submitted to specialists in the groups in

question, who will be requested to submit opinions on the nomenclatorial

status of said names.

(6) Upon return of the lists from the specialists, the Commission will

endeavour to test the names, according to the International Rules, and

if feasible will publish a list of the genera in question with their most

commonly used names and their correct names.

(7) If the undertaking is successful, the zoologists of the world will be in-

yited to give to the Commission the benefit of their criticisms not later

than July 1, 1912, so that the Commission can restudy the names and

submit to the next Congress :

(8) An official list of generic names, with their genotypes, and with the

(9) Proposition that the Congress adopt said list, and a

(10) Resolution to the effect that no zoologist shall upon NoMENCLATORIAL

grounds change any name in said list unless he first submits to the

Commission his reasons for making the change and unless the Com-

mission considers the reasons valid.

The Commission believes that this proposition is feasible, but

for the present views it in the light of an experiment, dependent

to no small extent upon the question whether a proper amount

of co-operation is forthcoming. In this connection the Commission

takes the liberty of inviting attention to the fact that the great

advances in nomenclature have been made by colleagues who have

showed a conviction in their view sufticient to induce them to

devote some time to the subject.

AMENDMENTS TO THE “ Régles internationales de la Nomenclature

zoologique.”’—In its executive sessions the Commission has eon-

sidered 30 propositions which have been submitted as amendments

to the present International Rules. Of these propositions, the

Commission unanimously recommends to the Congress the

adoption of the following :—

Art. 4. For the word root, substitute the word stem.

Art. 27 (6). For the word darva, substitute the words any stage in the life-

history.

Art, 35. Insert as a third paragraph the following :—

“Specific names of the same origin and meaning shall be considered

homonyms if they are distinguished from each other only by the following

differences ;

(a) The use of ae, oe, and e, as caeruleus, coeruleus, ceruleus; ei, i,and y, as

chiropus, chetropus; ¢c and k, as microdon, mikrodon.

(b) The aspiration or non-aspiration of a consonant, as oxyryncus, ory-

rhynchus.

(d) By a single or double consonant: ditoralis, littoralis.

(e) By the endings -ensés and -iensis to a geographical name, as timorensis,

temoriensis.”

Art. 36. Omit from the examples—Macrodon, Microdon ; caeruleus, coeruleus,

ceruleus ; silvestris, sylvestris, silvaticus, sylvaticus ; littoralis, litoralis; autum-

nalis, auctumnalis ; dana, danma.

Appendix F. In the English and German texts, substitute the words zrans-

literation and transliterated tor transcription and transcribed.

Appendix G. In all the three texts, substitute paragraph for rules, and omit

from the heading in French text the words [égles de la.

TTALIAN TRANSLATION.—The Commission has voted to issue an

official Italian edition of the International Rules.

(Signed) CH. WARDELL STILES,

Secretary of Connvission.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, February 7th, 1911, at half-past Hight

o'clock pP.M., when the following communications will be

made :—

1. Dr. W. N. F. Wooptanp, F.Z.S.

On the Structure and Function of the Gas-Glands and Retia

Mirabilia associated with the Gas-Bladder of some Teleostean

Fishes, with notes on the Teleost Pancreas.

2. Prof. J. Cossan Ewart, M.D., F.R.S., F.Z.S8.

Skulls of Oxen from the Roman Military Station at New-

stead, Melrose,

3. Groree P. Farran.

Plankton from Christmas Island, Indian Ocean.—I. On

Copepoda of the Family Coryezidee.

4, H. R. Hoge, M.A., F.ZS.

On some New Zealand Spiders,

The following communications have been received :—

1. Dr, F. D. Wetcsa, F.Z.S.

Observations on different Gibbons of the Genus Hylobates

now or recently living in the Society’s Gardens, and on a

Symphalangus syndactylus, with Notes on Skins in the Natural

History Museum.

2. R. LyDEKKER.

On Tragelaphus buxtoni, an Antelope from Abyssinia,

3. Epwarp G. BouLENGER.

a ee

A Contribution to the Study of the Variations of the Common

Salamander (Salamandra maculosa).

4. G, A. Boutencer, F.B.S., V.P.ZS.

On a Collection of Fishes from the Lake Ngami Basin,

Bechuanaland.

Communications intended for the Scientific Meetings should

be addressed to Sh * of

P. CHALMERS MITCHELL,

Secretary

ZOOLOGICAL Society oF Lonpon,

Recent’s Park, Lonpon, N.W.

December 20th, 1910.

MOTOR BUSES FOR THE SCIENTIFIC MEETINGS.

i On the night of each Scientific Meeting Motor ’Buses

B willrun direct to the Gardens from Regent’s Park Tube

F Station, starting at 8.5 and 8.20 p.m. The return

E *buses after the Meeting will leave the Gardens at 10.15

fand 10.35 p.m. These "buses will be labelled

/PRIVATE and no fare will be charged to Fellows

: attending the Meetings.

No. 90.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

February 7th, 1911.

Prof. E. A. Miycuty, M.A., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary read a Report on the additions that had been

made to the Society’s Menagerie during the month of December

1910.

Dr. C. W. Anprews, F.R.S., F.Z.8., exhibited a skull of a

Sabre-toothed Tiger (Smilodon californicus) from an asphalt

deposit in California, and pointed to anatomical characters which

tended to prove that the animal used its large canines for stabbing

and tearing, not for biting.

The Secretary exhibited a mounted specimen of the Platypus,

which had been lent for the purpose by Mr. P. St. Michael

Podmore, F.Z.8.

Mr. Epwarp GERRARD exhibited the head of a Caribou, shot

by Sir John Rogers, K.C.M.G., in British Columbia, which had

a distinct third antler over the centre of the orbital arch of the

frontal bone. There was a slight burr on a level with the skin,

but no pedicle. It had the appearance as if it could be shed in

the same manner as the normal antlers.

A very fine Eland head, obtained by Major Gordon on the

Bahr-el-Ghazel, and three fine heads of White-tailed Deer were

also exhibited, one pair of antlers being abnormal.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Sir Shillings per annum, payable in advance.

Dr. W. N. F. Wooptanp, F.Z.8., gave an account of a paper

on the structure and function of the gas-producing mechanism

(‘red body ”) found in connection with the gas-bladders of many

Teleostei (Physoclisti and Physostomi). After summarizing some

of the principal facts known concerning these subjects, the author

discussed various theories already advanced to account for the

details of gas-production, and showed that the most satisfactory

hypothesis was a combination of the views of Jaeger and of

Nusbaum and Reis, supplemented by additional facts and sug-

gestions then advanced in the paper. Among the additional

physiological facts adduced were the striking distribution of the

capillaries of the rete mirabile of the erythrocytolytic granules

resulting from the action of Jaeger’s toxin on the blood, and the

peculiar endothelium of the arterioles towards the proximal pole

of the rete. Both of these facts, together with the otherwise

meaningless conformation of the rete, supported the view already

advanced by the author that the rete mirabile duplex is essen-

tially a device to ensure the diffusion into the arterioles of the

toxin poured into the blood by the gas-gland, so that erythro-

cytolysis shall be well advanced (and therefore the abstraction of

oxygen made easy) by the time the arterial blood reaches the

gas-‘‘ secreting” cells. The author also made suggestions con-

cerning the exact meaning of hemolysis in connection with the

abstraction of oxygen from the blood by the gas-gland cells. The

first part of the paper dealt with the comparative morphology

of gas-glands.

Prof. J. Cossar Ewart, M.A., F.R.S., F.Z.8., gave an account

of his memoir entitled ‘‘Skulls of Oxen from the Roman Station

at Newstead, Melrose,” illustrating his remarks by lantern-slides.

He stated that examination of the skulis from Newstead lent

support neither to the descent of all Huropean cattle from the

Urus (Bos primigenius) nor to the descent of all European,

Indian, and African breeds from the Asiatic Urus (B. nomadicus).

He dealt with the evidence to be derived from the maxille, the

occiput and the temporal fosse, and stated his conclusions as

follows : ;

1. That the Celtic Shorthorn (Sos longifrons) is probably more

intimately related to the Zebu of India (Bos indicus) than

to the European Urus (Sos primigenius).

2. That long premaxille are usually correlated with an occiput

of the Bos primigenius type, while short premaxille are

usually correlated with an occiput of the Bos acutifrons

type.

3. That polled black Galloway cattle and polled white “ wild”

Cadzow cattle are intimately related to the Urus, that

flat-polled A berdeen- Angus cattle probably include amongst

their ancestors an ancient Oriental race now represented

by, amongst others, a Syrian breed with rudimentary

horns, and that round-polled cattle may belong to a still

more ancient Oriental race descended from Bos acutifrons

of the Punjab Siwaliks.

‘

€

Mr. G. P. Farran presented a paper, communicated by Dr. W.

T. Catman, F.Z.8., on Copepoda of the family Coryceide col-

lected by Sir John Murray, K.C.B., F.R.S., and Dr. C. W.

Andrews, F.R.S., F.Z.S., at Christmas Island. The collection,

though small in bulk, was exceedingly rich in species, and the

genus Coryceus was especially well represented.

A new genus was proposed and several new species were

described and figured.

Mr. H. R. Hoae, M.A., F.Z.S., read a paper on “Some New

Zealand Spiders,” based on a small collection sent by Prof. Charles

Chilton, of Christchurch, New Zealand. Twelve species and eleven

genera were represented in the collection, and a new local variety

of Tetragnatha ferox and four new species were described.

Mr. Otprietp Tuomas, F'.R.S., F.Z.S., read a paper on Mam-

mals collected in the Provinces of Kan-su and Sze-chwan, Western

China, by Mr. Malcolm Anderson, for the Duke of Bedford's

Exploration of Eastern Asia. This collection, from a region

hitherto almost unrepresented in the British Museum, was perhaps

the finest that had ever come from China, at least so far as small

mammals were concerned. 47 species were included, represented

by 350 specimens, presented, as on previous occasions, to the

National Museum by His Grace.

Besides examples of a number of rare and important species

not hitherto in the Museum, specimens of the following new

forms were included :—

PLECOTUS ARIEL, Sp. n.

Allied to P. wardi, but much darker in colour.

Forearm 44 mm.; thumb, without metacarpal, 9:2. Skull

17°72 x 9-4.

Hab. Ta-tsien-lu, Sze-chwan. Type. Female, No. 2343.

MyorTiIs ALTARIUM, sp. n.

Size large. Ears long and rather narrow. Skull with very

short muzzle.

Forearm 45 mm.; ear 22. Skull 15:2.

Hab. Omi-san, Sze-chwan. Type. Female. No. 2423.

SoREX BEDFORDIA, sp. n.

A small dark species, with a blackish line down the back.

Head and body 55 mm.; tail 55; hind foot 13. Skull 17-4 x 84,

Hab, Omi-san, Sze-chwan. Type. Male. No. 2541.

SOREX WARDI, sp. n.

Paler than S. bedfordie ; tail bicolor ; brain-case much smaller,

Head and body 53 mm.; tail 49; hind foot 12. Skull 17 x 8-2.

Hab. Tau-chow, Kansu. TZype. Male. No, 2280,

SORICULUS SACRATUS, sp. n.

Allied to S. caudatus, but brain-case smaller.

Head and body 60 mm. ; tail 54; hind foot 14. Skull 18:1 x 9-5.

Hab. Omi-san. Type. Female. No. 2485.

CHODSIGOA SMITHII, sp. n.

Proportions of tail as in Ch. hypsibia, but size much larger.

Head and body 85 mm.; tail 68; hind foot 18. Skull 22x

10:5.

Hab. Ta-tsien-lu. Z'ype. Male. No. 2349,

MUSTELA RUSSELLIANA, sp. Nn.

Size excessively small. Colour nearly as in UM. kathiah. Tail

not tufted.

Head and body 133 mm.; tail 54; hind foot 22. Skull 29°3 x

15:2.

Hab. Ta-tsien-lu. Type. Female. No. 2388.

EPIMYS EXCELSIOR, Sp. Nn.

Allied to #. confucianus, but larger. Skull longer and

slenderer.

Head and body 178 mm.; tail 193 ; hind foot 30. Skull 39.

Upper molars 6:8.

Hab. Ta-tsien-lu. Zype. Male. No, 2379.

EPpiMys ANDERSONI, sp. n.

Still larger than ZL. ewcelsior ; tail very long.

Head and body 164 mm. ; tail 248; hind foot 37. Skull 43.

Molars 8-2.

Hab. Omi-san, Sze-chwan. Type. Female. No, 2478.

APODEMUS FERGUSSONI, Sp. n.

General characters of A. agrariws, but no trace of a dorsal line.

Head and body 107 mm.; tail 98; hind foot 24; ear 15.

Skull 27-7.

Hab. Wen-hsien County, 8. Kansu. Type. Female. No. 2296.

PROEDROMYS BEDFORDI, g. & sp. nn. (Microtine).

Incisors grooved. M* with posterior lobe cylindrical, external.

Fur very long.

Head and body 103 mm.; tail 41; hind foot 18. Skull 26x 16.

Hab. 8.K. of Min-chow, Kansu. Type. Female. No. 2214.

Microtus (CARYOMYS) EVA, sp. n.

(Caryomys, subg. u.—Type, MZ. inex Thos.)

Allied to Mf. imez, but tail very much longer.

Head and body 88 mm.; tail 50; hind foot 16°5. Skull 22:-6x

13:5.

Hab. Near Tau-chow, Kansu. Jype. Male. No. 2238,

MIcROTUS MALCOLMI, sp. n.

A medium-sized species with high arched skull.

Head and body 103 mm. ; tail 40 ; hind foot 17. Skull 26-2 x 15.

Hab. 8.E. of Tau-chow, Kansu. Type. Male. No. 2245.

MICROTUS IRENE, Sp. 0.

Skull flattened; brain-case small in proportion to muzzle.

M, with only four closed spaces.

Head and body 98 mm.; tail 37; hind foot 18. Skull 24:2 x

14°8.

Hab. Ta-tsien-lu, Sze-chwan. Type. Female. No. 2396.

MYosPALAX CANSUS SHENSEIUS, subsp. n.

Colour warmer than in true cansus. Tooth-row decidedly

longer.

Head and body 178 mm.; tail 54; hind foot 30. Upper

tooth-row (alveoli) 11-9.

Hab. Yu-lin-fu, Shensi. Zype. Male. B.M. No. 9.1.1.216.

RHIZOMYS DAVIDI, sp. n.

Soft fur and short tail of 22. vestitus, but skull scarcely larger

than in &. sinensis.

Head and body 350 mm.; tail 69; hind foot 42. Skull 66 x

A9°5.

Hab. Kuatun, Fo-kien. Zype. Female. B.M. No. 96.12.1.6.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, February 21st, 1911, at half-past Hight

o'clock pP.m., when the following communications will be

made :—

1. Dr. H. G. Puen, F.R.S., F.Z.8., Pres.R.M.S.

Report on the Deaths which occurred in the Zoological

Gardens during 1910.

2. R. LYDEKEKER.

On Tragelaphus buxtoni, an Antelope from Abyssinia.

3. Epwarp G. BouLENGER.

A Contribution to the Study of the Variations of the Common

Salamander (Salamandra maculosa).

4. G. A. Boutenerr, F.R.S., V.P.Z.S.

oe oo eae

On a Collectlon of Fishes from the Lake Ngami Basin,

Bechuanaland.

5. Dr. F. D. Weucu, F.Z.S.

Observations on different Gibbons of the Genus Hylobates

now or recently Living in the Society’s Gardens, and on a

Symphalangus syndactylus, with Notes on Skins in the Natural

History Museum.

The following communications have been received :-—

1. R. LypEKKeEr.

(a) A rare Beaked Whale.

. (b) Age Phases of the Rorqual.

2. Mrs. E. W. Sexton.

On the Amphipod Genus Leptocheirus.

‘3. FV. BE. Bepparp, M.A., F.R.S., F.Z.S.

(a) Contrikutions to the Anatomy of the Anura.—I. Some

Anatomical Notes upon the Frog Megalophrys (Leptobrachium)

ee.

(6) On the Spermatophores in Earthworms of the Genus

Pheretima (= Pericheta).

4. Wiuitam Niconn, M.A., D.Sc., M.B.

On Three new Trematodes from Reptiles.

eC Sue TUN NUS IUaID Ne, ey OLA A.

Notes on Marine Ostracoda from Madeira. .

6. J. Lewis Bonnors, M.A., F.L8., F.Z.8, and F. W. Smaiey,

F.Z.S.

On Colour and Colour-pattern Inheritance in Pigeons.

7. P. CHALMERS Mrrenntn, NMEA Discs) Bonelli Dies.

F.ZS.

On Longevity and Relative Viability in Mammalsand Birds;

with a Note on the Theory of Longevity.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL SocreTY OF Lonpoy,

REGENT’s Park, Lonpon, N.W.

February 14th, 1911.

MOTOR ‘BUSES FOR THE SCIENTIFIC MEETINGS.

On the night of each Scientific Meeting Motor ’Buses

will run direct to the Gardens from Regent’s Park Tube

Station, starting at 8.5 and 8.20 p.m. The return

*buses after the Meeting will leave the Gardens at 10.15

and 10.35 p.m. These ’buses will be labelled

PRIVATE and no fare will be charged to Fellows

attending the Meetings.

No. 91.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

February 2ist, 1911.

Dr. A. Smita Woopwarp, F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary read a Report on the additions that had been

made to the Society's Menagerie during the month of January

OIE

Mr, A. Heneace Cocks, F.Z.8., exhibited a series of photo-

graphs of the female Brindled Gnu recently born in the Society’s

Gardens, and gave a brief account of its growth and coloration.

Dr. H. Hammonp Smiru, M.R.CS., F.Z.S., exhibited three

skins of male Pheasants assuming female plumage, sent to him

by Mr. Arthur Gilbey, and some microscopical specimens of the

glands prepared by Dr. S. G. Shattock.

Dr. H. G. Purmmer, F.R.S., F.Z.S., Pathologist to the Society,

reported on the Deaths which had occurred in the Zoological

Gardens during the past year, and illustrated his remarks with

a large series of lantern-slides, prepared from pathological material

obtained in the course of his work at the Gardens.

Mr. R. LyDEKKER communicated a short paper on Tragelaphus

buaxtoni, an Antelope obtained by Mr. Ivor Buxton in Abyssinia.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance.

Mr. EK. G. BouLENGER read a paper, communicated by Mr. G. A.

Bouncer, F.R.S., V.P.Z.8., on the varieties of the Spotted

Salamander (Salamandra maculosa), illustrated by lantern-slides

and specimens. One of the principal results of the author's

study was to lay greater stress on the disposition of the spots

than on their actual form, size, or colour, and to define two

principal forms in Central Europe, which had not previously

been separated with sufficient precision, notwithstanding their

well-marked geographical distribution.

The author further dealt with some of the experiments of

Dr. Kammerer, of Vienna, and the conclusions arrived at by him

with regard to coloration in relation to environment.

Mr. G. A. Bounencer, F.R.S., V.P.Z.S., contributed a paper

hased on a collection of Fishes from the Lake Ngami Basin,

Bechuanaland, made by Mr. R. B. Woosnam, F.Z.S.

This paper will appear in the ‘ Transactions’ In due course.

Dr. F. D, WetucH, F.Z.S., communicated a paper on Gibbons of

the Genus Hylobates, and on a Siamang Gibbon, recently living

in the Society’s Gardens, with notes on skins in the British

Museum (Nat. Hist.).

The next Meeting of the Society for Scientific Business will

be held on Tuesday, March 7th, 1911, at half-past Hight

o'clock p.m., when the following communications will be

made :—

LANTERN EXHIBITIONS :—

@. Habits of Moose.

b. Carl Hagenbeck’s Tierpark,

1. The Hon. N. Cuarues Roruscuiyp, M.A., F.Z.S.

Some new Siphonaptera from China.

2. F. E. Bepparp, M.A., F.RB.S., F.Z.5.

(a) Contributions to the Anatomy of the Anura.—I. Some

Anatomical Notes upon the Frog MJegalophrys (Leptobrachium)

fee.

(6) On the Spermatophores in Earthworms of the Genus

Pheretima (= Pericheta),

3. BR. LYDEKKER.

(a) A rare Beaked Whale.

() Age Phases of the Rorqual.

4, P. Cuaumers Mircuenn, M.A., D.Sc., Hon.LL.D., F.R.S.,

F.Z.S.

On Longevity and Relative Viability in Mammals and Birds;

with a Note on the Theory of Longevity.

The following communications have been received :—

1. Mrs. E. W. Sexton,

On the Amphipod Genus Leptocheirus.

2. Wiiiiam Nicoun, M.A., D.Sec., M.B.

On Three new Trematodes from Reptiles.

3. G. Stewarrson Brapy, M.D., LL.D., D.Sc., F.R.S., C.M.ZS.

Notes on Marine Marine Ostracoda from Madeira.

4. J. Lewis Poneotr, M.A., F.LS., F.Z.8., and F. W. Smairy,

E.Z.8.

On Colour and Colour-pattern Inheritance in Pigeons,

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary,

ZOOLOGICAL Society oF LonDoN,

Recgent’s Park, Lonpox, N.W.

February 28th, 1911.

MOTOR ‘BUSES FOR THE SCIENTIFIC MEETINGS.

On the night of each Srientific Meeting Motor ’Buses

will run direct to the Gardens from Regent’s Park Tube

Station, starting at 8.5 and 8.20 p.m. The return

‘buses after the Meeting will leave the Gardens at 10.30

and 10.45 p.m, These ’buses will be labelled

PRIVATE and no fare will be charged to Fellows

attending the Meetings.

No. 92.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

March 7th, 1911.

Dr, A. SmitH-Woopwarp, F.R.S., Vice-President,

in the Chair,

The Minutes of the last Scientific Meeting were confirmed,

Mr. R. JI. Pocock, F.L.S., F.Z.8., exhibited, on behalf of

Mr. E. C. OBERHOLTZER, a large number of lantern-slides and

photographs illustrating the habits of Moose. Mr. Oberholtzer

had spent several months in the forested lakeland tributary to

Rainy Lake, Ontario, photographing and observing the habits of

Moose, and had presented to the Society the series of enlarge-

ments exhibited.

The SrecreTary exhibited a series of lantern-slides prepared

from photographs kindly given to him by Mr. Cart HacEnsBeEck,

Silver Medallist of the Society, and illustrating some of the most

remarkable features of Mr. Hagenbeck’s new Tierpark at Stellingen

near Hamburg.

Dr. Karu Jorpan, F.E.S., and the Hon. N, Cuartes Roru-

SCHILD, M.A., F.Z.8., F.E.S., communicated a paper “On some

Siphonaptera from Northern China.” This collection of Fleas

had been made by Mr. M. P. Anderson, the Duke of Bedford’s

collector, in the province of Shen-si, and contained altogether 17

species, of which no less than 13 were new. Some of these

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in adyance.

were possibly only geographical developments, but others repre-

sented distinct types not very nearly allied to any known species.

Mr. F. EK. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the

Society, read a paper dealing with certain points in the anatomy

of the Frog Megalophrys (Leptobrachiwm) fec, based on specimens

which had been exhibited in the Society’s Gardens.

Mr. Brpparp also read a paper on the Spermatophores in

Harthworms of the genus Pheretima (=Pericheta). These

structures had been found in two species, one of which would be

described as new, contained in a collection of terrestrial Oligo-

cheeta from the Philippine Islands, which had been submitted to

him for examination by the Director of the Scientific Bureau of

the Philippines.

Mr. R. LypEKKER communicated two short papers entitled

(a) “A Rare Beaked Whale,” and (6) “Age Phases of the

Rorqual.”

Dr. P. Cuatmers Mircuery, M.A., F.R.S., Secretary of the

Society, gave an account of his memoir entitled ‘‘On Longevity and

relative Viability in Mammals and Birds; with a Note on the

Theory of Longevity.” The work was based ona study of the

records of the duration of life in the Society’s Gardens of over

20,000 individual Mammals and Birds. These were arranged

systematically so as to make possible a comparison of the average

duration with the maximum duration and what was known or could

be inferred as to the potential longevity. Such a method gave a

measure of the effect of the conditions of captivity on the duration

of life. The memoir discussed some of the results obtained by

such a comparison, particularly with regard to the provision of

artificial heat. In the note on the Theory of Longevity, the

author briefly reviewed the contributions of Ray Lankester, Weis-

mann, and Metchnikoff, and stated his conclusion that potential

longevity was due to constitutional causes, that the constitution

was adapted to the average specific longevity, and that the corre-

lation between longevity and reproduction was the reverse of

what had been suggested by Weismann.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, March 21st, 1911, at half-past Hight

oclock p.m, when the following communications will be

made :—

1. Mrs, E. W. Sexton.

On the Amphipod Genus Leptocheirus.

2. J. Lewis Bonnorn, M.A., F.L.S., F.Z.S., and F. W. Smauuey,

Se ee ee

HES:

On Colour and Colour-pattern Inheritance in Pigeons.

3. G. STEWARDSON Brapy, M.D., LL.D., D.Se., F.R.S., C.M.Z.S.

Notes on Marine Ostracoda from Madeira.

The following communications have been received ;-—

1. WiniiamM Nicout, M.A., D.Se., M.B.

On Three new Trematodes from Reptiles.

2. J. A. Morcu.

On the Natural History of Whalebone Whales.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Socrety oF Lonpon,

Recent’s Park, Lonpon, N.W.

March 14th, 1911.

MOTOR ‘BUSES FOR THE SCIENTIFIC MEETINGS.

| On the night of each Scientific Meeting Motor ’Buses

| will run direct to the Gardens from Regent’s Park Tube

F Station, starting at 8.5 and 8.20 p.m. The return

; buses after the Meeting will leave the Gardens at 10.30

Hand 10.45 p.m. These *buses will be labelled

f PRIVATE and no fare will be charged to Fellows

q attending the Meetings.

Exhibitions and Notices (continued).

Page

Mr. D. Seru-Surtu, F.Z.S. Exhibition of skins of the Australian Yellow-rumped Finch

Dr.

(Mairi flaipryInd): en esc ac oa ieee dis vee nals dime sie e/sleja ems aicinie a sels aes v's dale es 101

W. E. Hoyun, M.4., F.Z.S8. Explanation of the Report of the International Com-

mission on Zoological Nomenclature presented to the Graz Meeting of the International

WAMU LCA MUONS sha oreles waco nib, 6 sre, eyalla'n erdi'u Siem’ nie, Oe 6 nin. viprade sialalve' ein) erst aeavetelote old ollel 101

PAPERS.

1. On the Inheritance of the Webfoot Character in Pigeons. By J. Luwis Bonnore,

M.A., F.L.S., F.Z.S. (Text-figs. 3&4.) .......- 3 SUE eh CRE ar BAD KS 14

2. Notes on the little-known Lizard Lacerta jacksoni Blgr., with Special Reference to its

Cranial Characters. By Epwarp Drcen, F.Z.S. (Text-figs. 5-7.) ..........2.e00 19

3. On the Peloponnesian Lizard (Lacerta peloponnesiaca Bibr.). By G. A. BounEencnr,

HAR SUV eLEZeSeer (oleate sean cilllext ati oan) men atiponeianers teiarel ea -leh (aia! satolatohs cy 4 adeeat tats 37

4. Remarks on Two Species of Fishes of the Genus Gobius, from Observations made at

ROECOn. by; PD WARD, GS OUNMNGNOR Wat lrstolay qe 4 arnicla'sty's soe wielujers oi viaieel dele etc adeielnte 4)

5. On a Possible Cause of Pneumo-enteritis in the Red Grouse (Lagopus scoticus). By

H. B. Faytuam, D.Sc., B.A., F.Z.S., and H. Haumonp Situ, M.R.C.S., L.R.C.P.,

DENSZABS citar sal is) crass (oe ue (aheualy tener aac feted Seomsrtine oak char were stetals aletetniel > GieNate a: Srolarerareher cetera, seein 46

6. On the Alimentary Tract of certain Birds and on the Mesenteric Relations of the

Intestinal Loops. By Frank E. Bepparp, M.A,, ER.S., E.Z.8., Prosector to the

Rarenven | CMext=tins. OG) Momgmaiats ies, scadl's a.a'sibt oalalevetetens ciancieiatwre aie ake aa make AT

7. On the Specimens of Spotted Hyzenas in the British Museum (Natural History). By

Pro mPAN GED CABRIUAM CAMO! \'eo sass 5 ciel ga ostels mee onde a ele uy ate ile tial, vente 93

8, On the Segmentation of the Occipital Region of the Head in the Batrachia Urodela.

By Evwriy 8. Goopricu, M.A., F.R.S., F.Z.8., Fellow of Merton College, Oxford.

ReWeesctdt tem heUla es ages ase cots ahh a 6d a ay w aise ahe gr ogedete Cals cea eramenle oa Eka eks See Meee. Pe 101

9. The Mammals of the Tenth Edition of Linneus; an Attempt to fix the Types of the

Genera and the exact Bases and Localities of the Species. By Oxprimnp Tuomas,

EUSA Sh erssesieratele/ tavele ch auelin\e set syaretayt! sacar cuca errata apo tet aN ee cyt Skyy imide 120

10. The Duke of Bedford’s Zoological Exploration of Eastern Asia.— XIII. On Mammals

from the Provinces of Kan-su and Sze-chwan, Western China. By Oupririp Tomas,

GT HS le aby FS A hs Ns RASA SPECI AB ogee iocuei ei canes che TO a aM URN APNG Sei 158

Puate I,

Lacerta: pelsporme macnn We i ia 284) a page 37

- so that the complete reference i is now vB. %. Ss. 1911, p.

is as follows :— ai

Part 1 issued in March. — ‘ eS

ah eRe nale PACs cone Helig ee ac

A . ’ st a Me ee

ae De a aah ROR Nias tf September, ><.

{ i Se hae . va

sanctus December.

‘ a! 4 ‘ \

Mbasaeeainge? 1910, Part IY. (pp. 857-1083), were pul

\ ee lith, USIUR Rak anaes ey =

PROCEEDINGS

OF THE

ie os ce

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZQOLOGICAL SOCIETY

OF LONDON

1911.

PART II.

CONTAINING Pages 181 to 555, with 15 Puates

AND 93 TEXT-FIGURES.

JUNE 1911.

PRINTED FOR THE SOCIETY,

SOLD AT THEIR HOUSE IN REGENT’S PARK.

| | LONDON :

| MESSRS. LONGMANS, GREEN, AND CO., |

| | PATERNOSTER ROW. fi bo

| i ori! fi Sah i

a [Price Twelve Shillings. | co aa Sy

SEP 29% if

Ww Ye :

aki? 4 oa + ~ @ CUAL >

LIST OF CONTENTS.

1911, Parr IL. (pp. 181-555).

EXHIBITIONS AND NOTICES.

Tue Srcrerary. Report on the Additions to the Society’s Menagerie during the month of

December OL ve reys sie sae oss 3s ae sore) ois olor is eve eoteta etepmeneheeets choke a ani ettaast tends beret heases heat

Dr. C. W. Anprews, F.R.S., F.Z.8. Exhibition of the skull of a Sabre-toothed Tiger

OS RWDHOD GAN ORDIGIS) 55.66 ob cochoss ba op bo cuco nose So aA cdo guaUon od IOS 350009%

‘Tus Secretary, Exhibition of a mounted specimen of the Platypus (Ornithorhynchus

paradoxUus) ....++..6- ee Gre nme One ERT ane oon MAMA mGSo.! AL cing GE renee

‘Mr. Epwarp Gerrarp, F.Z.8. Exhibition of the heads of a Caribou (Rangifer tarandus

caribou), an Hland (Zauwrotragus derbianus gigas), and three White-tailed Deer

(Dorcelaphus americanus macrurus), (Text-fig. 52.) ..0ceee eee cree ee cece tenes

‘THE SECRETARY. Report on the Additions to the Society’s Menagerie during the month

of January 1911 }

Dr. H. Hamumonp Smiru, M.R.C.S., F.Z.S. Hxhibition of skins of male Pheasants assum-

ing female plumage ...

Mr. G. A. Boununeer, F.R.S., V.P.Z.S. Notice of a paper “On a Collection of Fishes

from the Lake Ngami Basin, Bechuanaland ”

ec eee se eter ee we HP OF oe oe ee OOP ewewesee

Mr. Aurrep H. Cocxs, M.A., F.Z.8. Exhibition of photographs of, and remarks upon, the

female Brindled Gnu recently born in the Society’s Gardens. (Text-figs. 97 & 98.) ..

‘Tue Secrutary. Exhibition of a series of lantern-slides illustrating some of the most

remarkable features of Mr. Carl Hagenbeck’s new Tierpark at Stellingen

ee ee ce tetas

Mr. Ervest ©. Osrrnonrzer. Exhibition of lantern-slides and photographs illustrating

an account of some Observations on Moose

PAPERS.

11. On the Structure and Function of the Gas Glands and Retia Mirabilia associated with

the Gas Bladder of some Teleostean Fishes, with Notes on the Teleost Pancreas. By

W.N. F. Woopnann, F.Z.8., The Zoological Department, University College, London.

(Plates II.-IX. and Text-figs. 53-62.)

12. On Skulls of Oxen from the Roman Military Station at Newstead, Melrose. By J.C.

Hwan, MD; BORIS. Geanse) (Bext-figs: Gaol) aj atein lett. felalc ieee ets eaeren telat ete atone

18. Plankton from Christmas Island, Indian Ocean.—I. On Copepoda of the Family

Coryexide. By Guorce P. Farran. (Plates XXIV.)

Ce rr nd

14, On some New Zealand Spiders. By H. R. Hoge, M.A., F.Z.S. (Text-figs. 92-96.) ..

Page

181

314

358

183

249

282

297

Contents continued on page 3 of Wrapper.

—

THE ZOOLOGICAL SOCIETY OF LONDON.

ee eee

Tars Society was founded in 1826 by Sir Sramrorp Rarrnzs

Mr. J. Sasrne, Mr. N. A. Viaors, and other eminent Naturalists,

for the advancement of Zoology and Animal Physiology, and for the

introduction of new and curious subjects of the Animal Kingdom,

and was incorporated by Royal Charter in 1829,

Patron.

HIS MAJESTY

THE KING.

COUNCIL.

HIS GRACE THE DUKE OF BEDFORD, K.G., President.

Tue Kart or Atramont, F.S.A.

Pror. J. Rost Braprorp, M.D.,

D.Sc., F.R.S., Vice-President,

Lr.-Cozr. Sire R. Haverock

Cuarurs, M.D., K.C.V.O.

Atrrep H. Cocxs, Ese., M.A.

Tue Rr. Hon. tHe Earn or

Cromer, P.C.,G.C.B., G.C.M.G.

F. D. Dawrrey Drewirr, Ese.,

M.A., M.D.

Caartes Drummonp, Ise.,

Treasurer.

Sir Epwarp Duranp, Br., C.B.

Freperick Gitterr, Kse., Vice-

President.

Srpney F’. Harwer, Ese., M.A.

Sc.D., F.R.S., Vice-President.

Str Water Rover Lawrencn,

Br GaChenn

Sir Epmunp G. Loprr, Br.

E. G. B. Meapre-Watpo, Esue.,

Vice-President.

P. Caatmers Mrrcnent, Esa.,

Me AC Dice De hase

Secretary.

W. R. Oetnvre-Grant, Esa.

Aprian D. W. Pottocg, Ese.

Avusyn Truvor-Barrysz,

M.A.

AntHony H. Wrnertep, Ese.

A.Smira Woopwarp,lse.,LL.D.,

F.R.S., Vice-President.

Henry Woopwarp, Esa., LL.D.,

E.R.S., Vice-President.

Esa.,

The Society consists of Fellows, and Honorary, Foreign, and

Corresponding Members, elected according to the By-Laws. It

carries out the objects of its foundation by means of the collection

of living animals, by its Library, and by its Scientific Publications.

The Office of the Society, where all communications should be

sent, addressed to ‘The Secretary,” is open from Ten till Five,

except on Saturdays, when it closes at Two p.m.

The Library, under the superintendence of Mr. F. H. Waterhouse,

is open daily at the above hours, except in September.

The Meetings of the Society for General Business are held on

the third Wednesday in every month of the year, except in

September and Octeber, at half-past Four o’clock p.m.

The Meetings for Scientific Business are held fortnightly on

Tuesdays, except in July, August, September, and December and

January, at half-past Hight o’clock p.m.

The Anniversary Meeting is held on the 29th. of April, or the

nearest convenient day, at Four P.M.

The Gardens are open daily from Nine o’clock until Sunset.

Mr. R. I. Pocock, F.R.S., F.L.S., is the resident Superintendent and

Curator of Mammals. Mr. D. Seth-Smith is Curator of Birds

and Inspector of Works. The Prosectorium for Anatomical and

Pathological work is under the charge of Mr. Frank H. Beddard,

M.A., F.R.S., Prosector, assisted by Mr. H. G. Phimmer, F.R.S.,

M.R.C.S., Pathologist to the Society.

TERMS FOR THE ADMISSION OF FELLOWS.

Frrrows pay an Admission Fee of £5, and an Annual Contri-

bution of £3, due on the Ist. of January, and payable in advance,

or a Composition of £45 in lieu thereof; the whole payment,

including the Admission Fee, being £00.

No person can hecome a Frrxow until the Admission Fee and

first Annual Subscription have been paid, or the annual payments

have been compounded for.

Frtiows elected after the 31st. of August are not liable for the

Subscription for the year in which they are elected.

———

a Le

PRIVILEGES OF FELLOWS.

Fettows haye Personal Admission to the Gardens upon signing

their names in the book at the entrance gate, and may introduce

‘Two Companions daily.

The Wire or Huszanp of a Futiow can exercise these privileges

in the absence of the Fellow.

Every Frtzow is entitled to receive annually 60 undated Green

Cards, and, when no specific instructions are received, the supply

will be sent in this form. If preferred, however, 20 Green Cards

may be exchanged for a book containing 2 Orders for each

Saturday * throughout the year. A similar book of Sunday Orders

may also be obtained in lieu of 20 Green Cards. A Green Card

may also be exchanged for 2 Buff Cards for the use of Children

under 12 years of age.

It is particularly requested that Fellows will sign every Ticket

before it goes out of their possession. Unsigned Tickets are not

available.

Green and Buff Tickets may be used on any day and in any year,

but in no case can two Children be admitted with one Adult’s

Ticket, or an Adult be admitted with two Children’s Tickets.

The annual supply of Tickets will be sent to each Frttow on the

1st. of January in every year, upon filling up and returning the form

of Standing Order supplied to Fellows.

Frtiows are not allowed to pass in friends on their written

order or on presentation of their visiting cards.

Frrtows are exempt from payment of the fee for Painting,

Sketching, and Photographing in the Society’s Gardens.

Frttows have the privilege of receiving the Society’s ordinary

Publications issued during the year upon payment of the additional

Subscription of One Guinea. This Subscription is due upon the

Ist. of January, and must be paid before the day of the Anniversary

Meeting, after which the privilege lapses. Frttows are likewise

entitled to purchase these Publications at 25 per cent. less than

the price charged to the public. A further reduction of 25 per

cent. is also made upon all purchases of Publications issued prior

to 1881, if above the value of Five Pounds.

Fettows also have the privilege of subscribing to the Annual

Volume of ‘The Zoological Record,’ which gives a list of the Works

and Publications relating to Zoology in each year, for the sum of

* The Saturday Orders are not available if the Fellow introduces friends

personally on that day.

One Pound Ten Shillings. Separate divisions of volumes 39 to

42 can also be supplied. Full particulars of these publications can

be had on application to the Secretary.

Fertows may obtain a Transrerabre Ivory Tickrr admitting

two persons, available throughout the whole period of Fellowship,

on payment of Ten Pounds in one sum. A second similar ticket

may be obtained on payment of a further sum of Twenty Pounds.

Any Frtiow who intends to be absent from the United Kingdom

during the space of at least one year, may, upon giving to the

Secretary notice in writing, have his or her name placed upon the

“dormant list,” and will then be called upon to pay an annual

subscription of £1 only during such absence, but after three years

must make a further application to be retained on that list.

Any Frttow, having paid all fees due to the Society, is at liberty

to withdraw his or her name upon giving notice in writing to the

Secretary.

Ladies or Gentlemen wishing to become Fellows of the Society

are requested to communicate with the undersigned.

P. CHALMERS MITCHELL,

Secretary.

Regent's Park, London, N.W.,

June, 1911.

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

FOR

SC ICIB IN, TUTE IP IEG) 15) WS IOINT JOS tsi

1911.

TurEspAy, OctoperR 24

" November 7 & 21

The Chair will be taken at half-past Eight o'clock im the Evening

precisely.

ZOOLOGICAL SOCIETY OF LONDON.

THE ZOOLOGICAL RECORD.

HE object of the Zootogicat Rucorpd is to give, by means of an

annual Volume, complete lists of the Works and Publications

relating to Zoology in all its branches that have appeared during

the year preceding the issue of the Volume, together with full

information as to the points they deal with, arranged in such a

manner as to serve as an Index to the literature of Zoology in all

parts of the globe, and thus to form a repertory that will retain its

value for the Student in future years.

The ‘ Zoological Record’ having been amalgamated with the

International Catalogue of Scientific Literature, Zoology, Volumes

from 43 onwards can now be obtained only from Messrs. Harrison

& Sons, except when purchasing complete sets from the Zoological

Society.

Under the scheme of amalgamation, Fellows of the Society, and

Institutions already on the subscription-list, have the privilege of

subscribing at the old rate of 30s. per annum, which covers the

cost of carriage of the volume. The subscription becomes due on

July Ist. in each year, and lapsesif not paid by the 1st. of December

following.

The Society is able to supply complete sets of the Record on the

following terms :—

Vols. 1 to 37, price £14 10s. net.

Vols. 38, 39, 40, and 41 at 10s. each net.

Vol. 42 and onwards at 40s. each.

The prices for separate volumes are as follows :—

Vols. 1 to 41 (except Vols. 4 and 6) 10s. each net.

Vol. 42 at 40s. The price of the ‘Zoological Record,’ Vol. 43 and

subsequent volumes, published now by Messrs. Harrison and Co.,

is 40s. each.

IypEx Zootoeicus. An alphabetical list of names of genera

and subgenera proposed for use in Zoology, as recorded in the

‘Zoological Record,’ 1880-1900; together with other names not

included in the ‘ Nomenclator Zoologicus’ of 8. H. Scudder. Com-

piled (for the Zoological Society of London) by Cuartzs Owen

Warernouss and edited by Davin Suarp, Editor of the ‘ Zoological

Record.’ London, 1902. Price to Fellows, 18s.; price to the

public, 20s., or if sold with a set, 10s.

Divisions of the ‘ Zoological Record’ of Vols. 39 to 42 can be

supplied by the Society, but after Vol. 42 they can be had only of

Messrs. Harrison & Sons, 46 St. Martin’s Lane, W.C.

[P. T. 0.

SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.

Divisions of the ‘ Zoological Record, Vols. 39-42, containing

the literature of the years 1902-1905, may be obtained separately

as follows :—

List of abbreviations of journals, etc.

Special Records, viz. :—

I. General Subjects ..

II. Mammalia

III. Aves ie eR

IV. Reptilia and Batrachia. .

V. Pisces Ree Cad

VI. Tunicata

VII. Mollusca

VIII. Brachiopoda ..

IX. Bryozoa

X. Crustacea

XI. Arachnida

XII. Myriopoda

XIII. Insecta

XIV. Kchinoderma

XY. Vermes ..

XVI. Coelenterata ..

XVII. Spongiz

XVIII. Protozoa

Hwwneo eR wd Fr £ pw dS & Oo bw

S 2G2EeQH eG2QG@Geeqeeqaeggsdoa¢g @

INS

Index of new names of genera and subgenera.

Divisions from Vol. 43 onwards are now supplied by Messrs.

Harrison & Sons, 46 St. Martin’s Lane, London, W.C.

P. CHALMERS MITCHELL,

Secretary.

Recent’s Park, Lonpon, N.W.

June, 1911.

ZOOLOGICAL SOCIETY OF LONDON.

LIST OF PUBLICATIONS.

Tue scientific publications of the Zoological Society of London

are of two kinds—“ Proceedings,” published in an octavo

form, and ‘ Transactions,” in quarto.

According to the present arrangements, the ‘‘ Proceedings”

contain not only notices of all business transacted at the scien-

tific meetings, but also all the papers read at such meetings

and recommended to be published in the ‘‘ Proceedings” by

the Committee of Publication. A large number of coloured

plates and engravings are issued in the volumes of the

“ Proceedings,” to illustrate the new or otherwise remark-

able species of animals described therein. Amongst such

illustrations, figures of the new or rare species acquired i im a

living state for “the Society” s Gardens are often given.

The “Pr oceedings”’ for each year are issued in four parts,

paged consecutively, on the first of the months of March,

June, September, and December. From January 1901 they

have been issued as two half-yearly volumes, indexed

separately.

An “ Abstract of the Proceedings ” is published by the

Society on the Tuesday following the date of Meeting to

which it refers. It is issued along with the < Proceedings,”

free of extra charge, to all Fellows who subscribe to the

Publications, but it may be obtained on the day of publii-

cation at the price of Sixpence, or, if desired, sent post free

for the sum of Six Shillings per annum, payable in advance.

The ‘‘ Transactions” contain such of the communications

made to the scientific meetings of the Society as, on account of

the nature of the plates required to illustrate them, are better

adapted for publication in the quarto form. They are issued

at irregular intervals.

Fellows and Corresponding Members, upon payment of

a Subscription of One Guinea before the day of the Anni-

versary Meeting in each year, are entitled to receive the

Society’s Publications for the year. They are likewise

entitled to purchase the Publications of the Society at 25 per

cent. less than the Bree charged for them to the Public. <A

further reduction of 25 per cent. is. made upon purchases of

Publications issued prior to 1881, if they exceed the value of

five pounds.

Fellows also have the privilege of subscribing to the

Annual Volume of the Zoological Record for a sum of 30s.

(which includes cost of delivery), payable on the Ist. of July

in each year; but tluis privilege is forfeited unless the

subscription be paid before the 1st. of December following.

The following is a complete list of the publications of the

Society already issued.

TRANSACTIONS* OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 19 vols. and Index. Ericete Jeol ip Hue

Fellows. Public.

Vol.. _I., containing 59 Plates.... (1833-85) .... £313 6 .... £418 OF

Ms il, +5 files veo (1835241) eee OO) SR oO ama

ays, UE - G3. yr ee ee S42 40) rea eo ato ue ASO

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Vol. XI., contaiming 97 Plates.. (1880-85) .... 912 0. IPAS

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PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND

CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF

LONDON. 8vo. 2 vols. (Letterpress only). Frice’ Exieeto ihe

Fellows. Public.

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yy Ul Ikea Ae es cree tnotons Ae OL 6 ASG. ate a lOse

PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.

8vo. 15 vols. (Letterpress only) and Index. (First Series.)

Price to Price to the Price to Price to the

Fellows. Public. Fellows. Public.

Tart J. 1883. 1 vol. 8vo. 4s. 6d. .. 6s.f | Part IX. 1841.1 vol. 8vo. 4s. 6d. .. 68.

, lies 4) faa on GS RM Cole Oman Iles. Ge.

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Wit 1080, 5 ARG oe Gap XV.1847. ,, 48, 6d. .. 6st

lu, WMO, 4 A Gah on Index 1830-1847, ,, 4s. 6d. .. 6s.

8vo. 138 vols. ‘tnd Index. (Second Series.)

Letterpress only. With Plates coloured.

Price to Price to the Price to Price to the

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Part MOWIL, IGAS, I all, Biro, 4 G2 50 ~CS% aa os cone Sl 0 8 s,s 7 Gy

eV MISO nae» 45.00) ve Reon eae 1 cOn38 4 le

Mar evs S50 ce, 4s, Gd) ou Ges eee 1.8 6.) isieanor

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XXVIII. 1860. . ASG) Wie (OSs. kaoaterieits Tne Ce ag er 2P Oe

Index 1848-1860. x As. 6d. 6s.

t Out of print.

* In consequence of a re-arrangement of the stock of the ‘Transactions,’ the Society is

now able to offer for sale, at the reduced price of £80, sets of Vols. v.—xvi. inclusive, and

separate papers, of which a list can be supplied, at about one-fourth their published price.

PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE

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BS Gime ASO take 13) GSE ox cae ec 9s. APs rh LOGS eres eat BH, Oth goog Cane

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a ae

ON ABNORMAL ANTLERS OF A CARIBOU. 181

EXHIBITIONS AND NOTICES.

February 7, 1911.

Prof. KE. A. Mincuin, M.A., Vice-President,

in the Chair.

THe Secrerary read the following report on the additions made

to the Society’s Menagerie during the month of December,

1910 :—

The registered additions to the Society’s Menagerie during

the month of December were 133 in number. Of these 76 were

acquired by presentation, 27 by purchase, 12 were received on

deposit, 17 in exchange, and 1 was born in the Gardens.

The total number of departures during the month, by death

and removals, was 216.

Amongst the additions special attention may be called to the

following :—

1 Viverrine Cat (felis viverrina), from India, 1 Northern Lynx

(Felis lynx isabellinus), from Tibet, 2 Binturongs (Arctictis bintu-

rong), from Malacca, and 1 Yellow-throated Marten (Mustela

Jlaviguia), from India, purchased on Dec. 24th; 4 Canadian

Skunks (Mephitis mephitica) and 2 Little Skunks (Spilogale

putorius), from N, America, the latter new to the Collection, pur-

chased on Dec. 29th.

1 Silver Fox (Vulpes ergentatus), from Hudson’s Bay, received

in exchange on Dec. 12th.

1 Feline Otter (Lutra jfelina), new to the Collection, from La

Plata, presented by William Clyne, Esq., on Dec. 6th.

1 Brindled Gnu (Connocheetes taurinus), born in the Menagerie

on Dee. Ist.

A collection of 15 Reptiles, including 1 Raddon’s Lizard (Wabuia

raddonit), 1 Smyth’s Snake (Grayia smythii), and 1 Yellow-

throated Snake (Zhrasops flavigularis), new to the Collection,

from Dunkwa, Gold Coast, presented by Dr. H. G. F. Spurrell

on Dec. 12th.

Dr. C. W. Anprews, F.R.S., F.Z.S., exhibited the skull of a

Sabre-toothed Tiger (Smilodon californicus), from an asphalt

deposit in California, and pointed to anatomical characters which

tended to prove that the animal used its large canines for stabbing

and tearing, not for biting.

Tue Secrerary exhibited a mounted specimen of the Platypus

(Ornithorhynchus paradoxus), which had been jent for the purpose

by Mr. P. St. Michael Podmore, F.Z.S.

Mr. Epwarp GERRARD, F.Z.S., exhibited the head of a Caribou

(Rangifer tarandus caribou) (text-fig. 52), shot by Sir John Rogers,

K.C.M.G., in British Columbia, which had a distinct forked third:

antler over the centre of the orbital arch of the frontal bone just

Proc. Zoou, Soc.—1911, No. XIII, 13

182 ON ABNORMAL ANTLERS OF A CARIBOU.

in front of the normal antler, which carried a single anterior basal

tine, and separated from it bya strip of hairy skin. There was a

slight burr on a level with the skin, but no pedicle. It had the.

appearance as if it could be shed in the same manner as the

normal antlers.

Text- fie. 52.

Head of a Caribou (Rangifer tarandus caribou) with abnormal antlers.

A very fine head of an Eland (Vaurotragus derbianus gigas) '

obtained by Major Gordon on the Bahr-el-Ghazel, and three fine -

heads of White-tailed Deer (Dorcelaphus americanus macrurus)

were also exhibited, one pair of antlers being abnormal.

it Sires,

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W.N.F.W. del.

ON THE GAS GLANDS OF SOME TELEOSTEAN FISHES. 183

PAPERS.

11. On the Structure and Function of the Gas Glands and

Retia Mirabilia associated with the Gas Bladder of some

Teleostean Fishes, with Notes on the Teleost Pancreas.

By W. N. F. Wooptayp, F.Z.S., The Zoological

Department, University College * adore

[Received November 9, 1916: Read February 7, 1911.]

(Plates II.-IX.t and Text-figures 53-62.)

ConvTENTs.

Page

Introductory .. SOE auE ES bcd oa dais -aRina ee Pee see Ree eon re Lees

Part [.—The Stauetaue of « ‘Red Bodies SAPs an Os iremetirs oe he hs URAL

Anguilla asa 'Type ...... 185

Other Examples of this First Type of “Red. Body’ My yr us, Ophichth. ys. 191

A Second Type of “ Red Body,” exemplified by Syngnathus.................. 193

Other Examples of this Second Type—G@obius, PRE CsOnEN. Ner nape:

Gasterosteus, Peristethus, Trigla, Smaris.. Dir. 195

Notes on the Teleost Pancreas, <7... Tee eee Ge LOS.

A Third Type of “ Red Body,” exemplified by Ophidium. BAB EEE pdec ecard 204

Other Examples of this Third Type—Bow, Atherina, Coris, Corvina,

Sargus, Balistes, Caranx, Serranus, Zeus, Gadus, Cepola, Perca . 205

Whe Classification of Gas Glands’ ..... gic 219

iheGiant Cellsior Gas) Glands) ro... erecta. 0 tine. 2g, 2

The Gas Bubbles in Gas Gland Cells ..............5, 196

Summary of Part I. and General Conclusions ‘respecting ‘Gas Gland

Structure... ‘ : BS Shin ic Saf ete aN bat pe

Part IJ.—The Physiolose of # “Red Bodies ” Loree astUn oh Qa

The Principal Theories as to the Origin of the Bladder Gases............... 290)

ANOS ccc boar PBQBEC Bebo ce nice Getic « =u COREE RES ERS eE SERENE roe EC.

Moreau.. Be is 3 SEER atte ste eth en Odean ays tein aacile aruavesle eegiem aoa eS!

Jaeger . Sire bebe cic on ccd pee NBR MRSS EPO R ALO OA ee oe ER PEAR Ae MRO?)

Nusbaum and Reis ......... PRLS Jn panne ne

Discussion of the Views of J: aeger ‘and Nusbaum & Reis. : 224

A Statement of the Modern View of the Physiology of the “Red

Bodies 777 secs 228

Some additional Suggestions by. the present Writer—the ‘Rationale of

the Rete Mirabile, the Disintegration of the Red Blood Cor ee

and other Minor Features of “ Red Body” Structure.. 230

ApprEnpiIx A.—The Sources and Modes of Preparation of Material. JSS oad ea baT 236

Apprnprx B.—On the Artificial Production of Gas Bubbles in Cells of the

Gas Gland |........-: : 237

IN cleric cts ceemrtetcs hes ON ets MAN ee peta ee nina ee cnet nae Mp asrer it FNC 3 241

Literature Lists . REN TAH RNS fi tem on eNe CUS SR PUES Rt! Siete BRIE Eo ee ET Fe

Iyrropucrory.

In the present paper it is proposed tu give a fairly complete

account of the gross and _ histological ereachare of the vasculo-

glandular organs or “‘red bodies” found in the wall of the gas

bladder = of many teleost fishes, and to offer certain suggestions

* The greater part of the work connected with the present paper was carried out

in the Zoological Department of King’s College, London.

+ For explanation of the Plates see pp. 245-248.

~ The terms “swim-bladder ” and “ aiv-bladder ” are inappropriate, since fishes do

not use this apparatus for swimming and it dees not contain-air,

Lor

184 DR. W. N. F. WOODLAND ON THE

respecting the physiology of these organs. Though the litera-

ture * concerned with the morphology and physiology of the gas

bladder is, of course, very considerable, yet up to the present only

one author (64, @5) has attempted a complete review of the

subject of the present paper in the light of recent work. Further,

although there exist several memows dealing with special types of

“ved body,” yet most of these are so occupied with cytological

and other details that, in the opinion of the present writer, they

have failed to supply such an account and such illustrations of

the gross structure of these “red bodies” as to render evident

some of their most remarkable features. Finally, the Jaeger-

Nusbaum controversy calls for an independent description of these

unique organs, so interesting from the physiological standpoint.

i have divided the present paper into two parts, the first of

- which deals with the structure and the second with the physio-

logy of the “red bodies.” With reference to the drawings

illustrating Part I., I have, with a few exceptions, adopted a

definite scheme of coloration in order to facilitate comprehension.

Green has been adopted to denote the internal lining epithelium

of the bladder, and therefore the glandular, 7. e. gas gland, epi-

thelium is always of this colour in the drawings, since this is but

a special development of the usually squamous lining epithelium ;

red denotes blood; black or grey has been employed for connec-

tive and muscular tissue and secretion material ; and yellow bas

been used to indicate the pancreas, which is often closely

associated in a mechanical sense with some parts of the “red

body.” The sources and methods of preparation of my material

are stated in Appendix A (p. 236).

Part I.—THE Srrucrure or THE ‘“ Rep Bopres” IN

vARIOUS TELEOSTET.

It is necessary to recognize at the outset that the term “red

body ” unfortunately has been apphed to different structures by

no means equivalent to each other: thus, e. g., the “ red bodies ” of

the Common Kel are simply the two retia mirabilia, whereas the

“red body” of the Cod consists both of numerous small retia

mirabilia and of the large mass of epithelial cells called the gas

gland, the retia and the gas gland being, in this case, intimately

associated. A gas gland exists in the Eel as in all other fish

with “red bodies,” but it is not in close connection with the

rete mirabile as inthe Cod. This being the case, it must be

understood that by the term “red body” all that is meant is a

ved mass situated in the bladder-wall, the constitution of which

is variable. I may add that in the vast majority of cases ‘“ red

body” includes both rete mirabile and gas gland. As an intro-

duction to the study of gas gland structure I shall first describe

the anatomy of one of the most simple as well as most common

* For the literatnreand historical reviews of the entire subject of gas bladders up

to 1866 sce Milne-Edwards (49) and Gouriet (84); for more recent literature see

the lists supplied by Bridge (22), Hiifner (38), Jaeger (44), Deineka (29), Reis &

Nusbaum (62, 63), and at the end of the present paper. ‘ ea

GAS GLANDS OF SOME TELEOSTEAN FISHES. 185

forms, viz. that of the Common Eel, Anguilla vulgaris. The “red

body” of the Hel has been described: several times previously,

e. g. by Jacobs (42, 43), Quekett (61), and Corning (28), but never,

J venture to think, in a sufticiently clear or comprehensive

manner and certainly never adequately illustrated.

The Vasculo-glandular Structure in the Bladder of the Common Hel

(Anguilla vulgaris).

It is well known that the bladder of the Eel is an elongated

sac lying dorsally and towards the posterior end of the body

cavity and connected with and opening into the esophagus by

means of a long wide duct—the ductus pneumaticus (text-fig. 53,

P.D., p. 186). The pneumatic duct opens into the bladder at about

midway i in its length, the bladder in consequence being divisible

into anterior (A. B. ) and posterior (P.B.) regions. The bladder

and duct are both situated in the same hor zontal plane, 7. e. the

duct opens into the bladder at the side, not dorsally or ventrally.

Running along the ventral side of the duct are an artery and

a vein situated close together (A.V.). This artery and vein give

rise to the two ovoid retia mirabilia (R.M.) where the duct opens

into the bladder, the two retia lying one on each side of the

duct and therefore being dorsal and ventral in position respec-

tively and in the same vertical line.

The wall of the bladder (see Pl. IT. figs. 1-6, e.g.) in fish generally

is best described as consisting of the tunica externa and tunica

interna. The former consists of two definite layers of fibrous tissue,

the outer being loose in texture, often containing muscular tissue

and varying greatly in thickness, the inner dense, silvery in appear-

ance, and principally consisting of elastic fibres. The latter consists

of a thin basis of dense connective tissue, with which is closely

connected the innermost layer of the bladder wail, a layer con-

sisting usually of flattened cells. It is this innermost cellular layer

which gives rise to the mass of glandular cells in connection

with a rete mirabile—the gas gland. The tunica interna is

easily separable from the tunica externa, and it is advisable to

remove this latter before cutting sections of the gland.

The vascular supply of the bladder may first be considered. It

has already been mentioned that an artery and a vein run along

the ventral side of the wall of the ductus pneumaticus. These

two vessels are situated in the outer fibrous layer of the tunica

externa (Pl. IT. fig. 1), which is here of considerable thickness, and

they supply several arteries to the duct and receive several veins

from it. Shortly before these two vessels reach the region of the

junction of the duct with the bladder, the artery and the vein

each divide into two (fig. 2), so that there are two pairs of vessels

instead of one, each pair consisting of an artery and a vein.

Now the artery and vein of each of fhese pairs undergoes further

subdivision, the smaller arteries and veins resulting from this

clic nero intermingling in their course, but noun once con

municating with each other, so that we now have situated towards

186 DR. W. N. F. WOODLAND ON THE

the upper and lower sides of the ductus pneumaticus two distinct

masses of intermingled arteries and veins (fig. 3), each mass

having resulted from the subdivision of the two pairs of arteries

and veins, which, as we have seen, have themselves originated

Text-fig. 53.

The bladder, duct, and “‘red bodies” of Anguilla vulgaris,

viewed from the side (after Quekett).

P.D., pneumatic duct which opens anteriorly into the esophagus; A and V, artery

and vein supplying bladder; R.M., rete mirabile; A.B., anterior 1egion of

bladder; P.B., posterior region of bladder.

i i

GAS GLANDS OF SOME TELEOSTEAN FISHES. 187

by the division of the initial artery and vein. The subdivision

of the arteries and veins in each mass proceeds rapidly, until

there is formed by this process an ovoid mass of extremely fine

parallel arterial and venous capillaries, closely intermingled with

each other, but never intercommunicating and indistinguishable

structurally as arteries and veins under the highest powers of

the microscope (fig. 4). These two ovoid masses of parallel

arterial and venous capillaries lying on the dorsal and ventral

sides of the ductus pneumaticus just anterior to its junction

with the bladder are the retia mirabilia of the eel-bladder (text-

fig, 53, R.M.).

The finest arterial and venous capillaries of each rete mirabile

run parallel to each other for a short distance, but soon they

commence to unite, arterial capillaries with arterial capillaries

and venous with venous, in order to re-form larger vessels similar

to those which produced the capillaries by subdivision. This

process of reunion proceeds so far as to form two masses of

large intermingled arteries and veins similar to those depicted

External aspect of a rete mirabile (after Quekett). The anastomoses of the

vessels shown in this figure do not occur in actuality.

in figure 3 (cf. figs. 3 & 5). When reunion has proceeded thus

far, each of these masses of re-formed arteries and veins once more

breaks up into fine capillaries in connection with the vascular

supply of the glandular folded epithelium which lines the bladder

internally. I reproduce here as faithfully as I am able (text-

fig. 54) Quekett’s figure (61) of the construction of the Hel’s rete

mirabile, which illustrates in a general way that which I have

just described. Text-fig. 54 is unsatisfactory in that it does not

indicate the fact that the vessels and capillaries forming the

rete are of two kinds—arterial and venous-—closely intermingled

with each other, although never intercommunicating ; also anas-

tomoses never occur between adjacent large vessels in the manner

shown in the figure. A more satisfactory diagram of the

construction of the Eel’s rete mirabile is the one I have

188 DR. W. N. F. WOODLAND ON THE

designed for text-fig. 55. In this the artery and the vein

giving rise toa rete mirabile are distinguished from each other

by the latter being cross-striped, the subdivision of each of

them is indicated, also the intermingling of the fine parallel

capillaries, their reunion and the second breaking up mto capil-

laries of the large arteries and veins thus formed to supply the

epithelium of the gas gland.

Up to the present I have spoken of the arterial and venous

capillaries or vessels composing each of the two retia mirabilia

merely as tubes, but if, of course, we take into account the

direction of the flow of blood in the two sets of vessels, then

it is evidently incorrect to speak of the arteries and veins both

subdividing at the anterior pole of the rete mirabile to produce

Text-fig. 55.

The construction of the rete mirabile bipolare geminum (R.M.).

G.H., glandular epithelium of gas gland.

capillaries: what is really happening in this region is that the

arteries are subdividing and the veins uniting (text-fig. 55).

That this is the actual mode of construction of the rete miabile

is not only proved by the study of serial sections but also by the

injection method. And here let me remark upon the extra-

ordinary process implied by this construction of the rete

mirabile—a term which, like its German rendering, ‘ wunder-

netze,” happens to be peeuliarly appropriate*. It is implied

that parallel vessels carrying blocd in opposite directions can so

influence each other, or be so influenced by a common cause, as

to break up into a rete mirabile at the same spot, the subdivision

of each of the two sets of vessels being so adapted to the other

as to ensure the close apposition and intimate intermingling

of their finest capillaries t. It is evident that this intricate

* Miiller (53) termed this type of rete mirabile a “ rete mirabile bipolare geminum, ”’

in contradistinction to a “rete mirabile bipolare simplex,” such as the purely arterial

carotid gland ot Amphibia.

+ The fact illustrated by these retia that blood-vessels are able to capillarize on

their own account, 7. e. quite apart from any other tissue, first attracted my attention

to these “ red bodies”? when engaged in certain speculations concerning the “renal-

portal system” in Vertebrates (76, 77). The formation of the retia seems to lend

support to Dr. Shore’s views (68, 69) concerning the vascularization of the liver

as distinguished from the venous penetration of the kidney-substance to form the

‘“renal-portal system.” I hope shortly to investigate the subjects of the “renal-

portal system” and the various kinds of retia mirabilia from a practieal standpoint.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 189

mechanism must be for some definite purpose, especially since it

is found in all fishes possessing gas glands. What that purpose

may be is a question I shall discuss later (see Part IT.).

The epithelium lining the ductus pneumaticus, which is sup-

plied with a few branches direct from the bladder artery and is

therefore not supplied by the vessels connected with the retia

mirabilia, is quite simple in character. It merely consists of a

single layer of flattened cells raised up at intervals by the large

capillaries running in the basis of connective tissue underlying

the epithelium (PI. II. fig. 7). These capillaries in the wall of the

duct apparently have much the same surface arrangement as those

found in the lungs of certain reptiles (Quekett, 61), but it is

hard to suppose from this fact alone that the epithelium of the

duct is normally respiratory in function, since the blood supplied

to this has already been oxygenated in the gills; on the other

hand, it is equally improbable that the arterial blood in the wall

of the duct serves to alter the composition of the adjacent gas

since there is reason to suppose that the duct normally serves as

a mere exit for the superfluous gas of the bladder (Jaeger, 45).

It is possible, however, that this large vascular supply of the duct

epithelium in the Eel is really correlated with the terrestrial

habits of the animal, since when an Hel travels across a meadow

the gillsare useless for obtaining oxygen, and under these circum-

stances the animal doubtless draws upon the oxygen in the

bladder—the duct acts as alung. Moreau proved that Perch,

when placed in water previously boiled so as to deprive it of

dissolved air, utilized the bladder-oxygen for purposes of respira-

tion. An interesting confirmation of this view is that in other

physostomes (Wyrus, Ophichthys, Hsox) the duct epithelium is, as

we shall see, practically devoid of capillaries *.

~ The lining epithelium of the bladder proper is very different

in character from that of the duct. It consists of a more or less

folded single layer of cells which are large in size and short

columnar in form, slightly granular, and, when stained by the

picro-indigo-carmine method (Appendix A), usually rather opaque

(PIDi. fie. 8). The folding of this layer is due to the presence of

large capillaries which push1 their way inwards towards the bladder-

cay ‘ity from the connective tissue base, and in so doing cause the epi-

thelial layer to form corresponding emergences. Thus the interior

of each projection of the glandular epithelium into the bladder

cavity is occupied by an extension of connective tissue enclosing

capillaries. The spaces situated between such emergences adjacent

to each other are also regarded as forming gland ducts. The

folding of the epithelium is very variable in different parts of the

bladder wall, in some parts almost disappearing and in others

being more accentuated than that depicted in figure 8; this

folding is much more marked in the anterior region of the

bladder than in the posterior. It might appear to be significant

that immediately the lining epithelium of the duct becomes sup-

plied with blood connected with the rete mirabile system of vessels

* JT find that Jacobs (42, 48) states this view as a fact.

190 DR. W. N. F. WOODLAND ON THE

(that is, when the duct opens into the bladder) it then assumes

the glandular folded character of the bladder epithelium, were it

not for the fact that in other physostomous genera this sharp

distinction between duct- and bladder-epithelium is absent, as I

shall show shortly.

The intracellular structures described in detail below as being

present in the cells of the gas glands of other teleost fishes—

intracellular gas bubbles, capillaries, and ducts—are not easily to

be detected in my preparations of the Hel, but I believe I have

seen gas bubbles present in a few cases, also the capillaries

occasionally intrude somewhat upon the cells at their bases.

Granular matter, ejected into the bladder-cavity with the gas, is

so small in amount in the gland ducts of my preparations that it

may be said to be absent.

The folded glandular epithelium of the gas gland of the Eel,

and presumably of all other genera possessing a folded epithelium,

originates from a singie unfolded layer which during development

becomes pushed out into the folds by the activity of the large

capillaries situated in the thin layer of connective tissue- lying

at the base of the epithelium *.

Thus in the type of “red body” found in the Common Hel

we have two large retia mirabilia, situated some distance from

the epithelium of the bladder which they supply, and the greater

part of this lining epithelium of the bladder is glandular and

folded in a simple manner to form the gas gland. In all “red

bodies ” we have these two elements, the rete mirabile and the gas

gland, but in the majority of ‘‘ red bodies” these two elements

are much more intimately associated with each other and more °

complex in form as compared with the structure of the simple type

of “red body” just described. The Hel type of “red body” serves

as a convenient starting-point for considering more complex types.

Before considering these types, it 1s necessary to point out a

serious mistake contained in almost the only English paper

dealing with the present subject, that of Vincent and Barnes (75),

which, though the paper is now quite out of date, has found its

way into at least one recent text-book (‘Cambridge Natural

History,’ vol. vii. 1904, p. 308). Vincent and Barnes, following

Coggi (25), state that “red glands are only found in those

bladders which are devoid of a ductus pneumaticus,” and they

draw a sharp distinction between Physoclisti, which possess the

gas gland, and Physostomi, which are supposed to be devoid of

one. I need hardly say that no such distinction exists. Vincent

and Barnes are right, in so far as they say that the ‘‘ red bodies ”

of the Hel are only partially comparable with those of the

Gadide (vide supra), but it is difficult to understand how they

came to overlook the large and conspicuous glandular epithelium

lining the bladder cavity. Further, as Deineka (29) points out, in

the physostomous Pike (Hsox luciws) a conspicuous gas gland

* J am much indebted to Dr. W. G. Ridewood for kindly presenting me with

most of my young material.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 191

exists (described by Coggi, 26), comparable in all essential respects

with that of the Cod. Also Vincent and Barnes, unlike many of

their predecessors, altogether fail to appreciate the peculiarity of

the arterial and venous nature of the rete mirabile; indeed, they

even appear to have some doubt as to the fact itself. They say :

“ Corning has discriminated between arteries and veins in what

we have usually spoken of as the ‘capillary masses.’ This

he has done by means of a series of injections. ... . This method

would certainly give some idea of the distinction between the

smallest arterioles and smallest venules, but we are doubtful

whether it would be more than a very rough method of distine-

tion, that it would be, in any given case, an infallible test as to

whether a particular small vessel were to be called arter y or vein.

Much would depend, it appears to us, on the strength of the

injecting force [!]. However this may be, we have failed to make

out anything in the minute structure which would warrant us in

dividing up the ‘ capillary masses’ into arterioles and venules.”

This statement suggests that Vincent and Barnes never worked

through a series of sections of the Eel (or any other similar)

“red body” (which possibly also accounts for their missing the

glandular epithelium of the Eel, which, as above stated, lies

separate from and posterior to the retia mirabilia *), since the

arterial and venous nature of the rete mirabilia is rendered quite

evident by this means alone; also the last-quoted statement of

Vincent and Barnes is still less comprehensible when we remember

that these “red body” retia mirabilia have been correctly

described in full and compared with the several other kinds

of retia mirabilia which exist by Johannes Miiller (53) in 1840,

not to mention Owen’s description (not confirmed by his figure,

however) of the ‘‘ red body ” of the Hel just referred to T.

Vasculo-glandular Structures of the Anguilla or First Type in the

ladders of some other Anguilliformes.

Myrus vuuearis Kaup.

In this physostome the vascular and glandular conditions are

similar to those found in Anguilla vulgaris. The only essential

distinction between the two is in the character of the lining

epithelia of the duct and bladder, and in this particular the

* A fact clearly stated by Owen (‘The Comparative Anatomy and Physiology of

Vertebrates, vol. i. 1866, p. 495): “The two chief ‘retia mirabilia’ or vaso-

ganglions, in the air-bladder of the Eel and Conger, which are situated at the sides

of the opening of the air-duct, are also‘ bipolar’ and consist of both arterioles and

venules [curiously enough, however, in contradiction of this statement, the figure

329 of the ‘ vaso-ganglion’ of the Eel supplied on p. 496 shows the blood going

through all capillaries in one direction !] : they consist of straight parallel capillaries,

as in fig. 329; their afferent trunks do not ramify in the immediate margin of the

vaso-ganglion "from which they issue, as in the vaso-ganglions of the Cod, Burbot,

Acerine, and Perch, but run for some distance before they again branch to form the

common capillary system of the lining membrane of the air-bladder.”’

+ These criticisms must not be held to imply any lack of appreciation of a very

lucid and interesting paper ; the fact, however, that the statements criticised have

been made use of by the late Professor Bridge so recently as 1904 shows the need for

correction.

192 DR. W. N. F. WOODLAND ON THE

distinction is striking. In Anguilla the epithelia of the duct and

bladder are, as we have seen, quite different in character; in

Myrus vulgaris, on the other hand, they are very similar. ‘The

duct epithelium consists of a single layer of short columnar cells

which is only slightly folded. Very few capillaries are present in

the connective tissue layer underlying the epithelium, though

here and there large capillaries are to be seen in transverse

section. Thus the duct is not lined by squamous epithelium as

in Anguilla. In the bladder of Myrus vulgaris the epithelium is,

as just stated, similar to that of the duct, the only difference

being that the cells are larger. The folding of the epithelium is

at most very slight and most pronounced in the region of the

opening of the duct; in the anterior and posterior regions of

the bladder, folding of the epithelium is practically absent. There

is never any approach to the degree of folding seenin Anguwilla.

It is also noticeable that where the epithelium passes over a large

capillary (and capillaries are not very numerous) the cells

immediately assume a more squamous character.

OPHICHTHYS (SPHHGEBRANCHUS) IMBERBIS Delar.

In this physostome the plan of construction of the “red

body ” is essentially the same as that found in Anguilla and Myrus,

but there exists in the single specimen at my disposal one

modification in connection with the rete mirabile. When

the bladder is viewed in situ, a single “red body” is seen to

be present which is situated anteriorly in the ventral wall where

the duct joins the bladder. A series of transverse sections shows

that this “red body” is, as in the Hel, a large rete mirabile,

which, however, is disposed in a different manner in the present

instance. The artery and vein run along the right side of the

bladder and break up to form the rete in such a manner

that the long axis of this lies transversely to the length of

the fish (Pl. I. fig. 9). The capillaries of the rete mirabile

unite as usual to form larger vessels, which here, of course, lie to

the left side of the bladder, where they break up anew to supply

the epithelium of the gas gland. As in Myrus, there exists no

sharp distinction between the glandular epithelium of the duct

and that of the bladder, also the capillaries in connection with

the epithelia are comparatively few in number. This epithelium

consists, as in the two preceding genera, of a single layer of

more or less columnar cells which occasionally becomes very

slightly folded, but over the greater part of the wall, both of the

bladder and the duct, the epithelium is a simple unfolded layer of

columnar glandular cells. Strange to say, the cells of the epi-

thelium of the bladder in my preparations are distinctly smaller

than those of the duct. Occasionally large capillaries are present

in the bladder wall, and where these oceur the epithelial cells

become, as in A/yrus, almost squamous in shape. Small capillaries

also exist, though they are not numerous. The epithelium of the

duct remote from the bladder becomes much more squamous in

character than elsewhere.

ee oe ee Pe ee

GAS GLANDS OF SOME TELEOSTEAN FISHES. 193

OPHICHTHYS (OPHISURUS) SERPENS L.

The general plan of the ¢wo retia mirabilia and the gas gland

is exactly that found in Anguilla. The epithelial lining is, as

in Anguilla, squamous and covers projections of the subjacent

connective tissue, which contain, not one large capillary, but many

small ones. Posteriorly the lumen of the duct, which anteriorly

is circular in transverse section, becomes very restricted and star-

shaped in outline, owing to the great size of the connective tissue

projections just mentioned—a feature not met with in the species

described above. The epithelium of the bladder wall is hardly

folded at all over the greater part of its area, simply consisting

of a single layer of short columnar glandular cells, but in places

folding occurs, occasionally to such an extent as to resemble

the bladder epithelium of Angwilla, but this is rare.

It is curious how the epithelial linings of the duct and bladder

differ in these four closely related fish—A nguilla, Myrus, Ophichthys

imberbis, and O. serpens. In the duct and bladder of Anguilla we

have the extremes of flattened and folded epithelia ; in the other

genera intermediate conditions of the epithelium exist, both as

regards the form of the cells and their arrangement, and the duct

and bladder epithelia are not sharply distinguishable from each

other, and yet the vascular supply has the same distribution in all

cases, although it undoubtedly varies considerably in amount,

being much greater in the Hel than in the other genera. The

different conditions of the rete mirabile in the two species of

Ophicthys are also noticeable.

The type of vasculo-glandular apparatus found in the bladders

of the preceding genera may be defined as follows: the glandular

epithelium is composed of a single layer of cells which either

remains unfolded or is only simply folded, and the retia mirabilia

are one or two in number and are large and distinctly separate

from the gas gland, the capillaries of the rete reuniting to form

large arteries and veins before coming into connection with the

glandular epithelium.

A Second Type of Bladder Vasculo-glandular Apparatus,

as exemplified i Syngnathus acus JJich,

Syngnathus acus is a physoclistous fish in possession of a “red

body,” which is, on the whole, very similar in construction to that

of the Eel, but which nevertheless exhibits certain differences

which justify us in separating it as a distinct type. The “red

body ” in Syngnathus is visible with the naked eye as a small

red patch at the extreme anterior end of the bladder. An artery

and vein running in the anterior attachment of the bladder divide

up in the manner already described for the Kel to form the single

rete mirabile (Pls. 11. G III. figs. 10-14). It willalso be observed in

figures 10 and 11 that posterior extensions of the diffuse pancreas

(indicated in the figures by yellow masses) penetrate into the

anterior end of the rete mirabile and, as will be shown in detail

194 DR. W. N. F. WOODLAND ON THE

later, are closely associated with the larger veins. Lying along-

side the rete mirabile is the anterior extremity of the elongated

closed bladder (fig. 14), the wall of the anterior region of which

is very glandular and vascular. A little way behind the anterior

extremity of the bladder, the side of the rete next the bladder

comes into connection with the bladder epithelium (fig. 15), the

capillaries uniting (¢. e. the arterial capillaries are uniting and

the venous capillaries are really here subdividing to form the

minute venous capillaries of the rete) to form slightly larger

vessels before supplying the epithelium of the gas gland. This

process of supplying the glandular epithelium goes on until the

whole of the rete mirabile is used up (PI. III. figs. 15-17,), the

fine capillaries of the rete always uniting to some extent to form

larger vessels before these supply the epithelium. ‘The bladder

epithelium, as shown in the figures, becomes enormously thickened

by a process of folding during the breaking-up of the rete mira-

bile. Posterior to the rete mirabile, which has thus disappeared

in supplying the glandular epithelium, this latter itself gradually

dwindles (fig. 18) until in the posterior region of the bladder

the cavity is lined with simple squamous non-glandular cells.

The epithelium of the bladder is quite simple in type and

essentially resembles the folded epithelium of the Kel, though

the folding is much more pronounced. Figure 19 represents the

glandular epithelium in the unthickened region, that is, lining

any side of the bladder cavity other than that adjacent to the

rete mirabile, where, as just stated, the epithelium is much

thickened. The folds of the epithelium due to the large capil-

laries which push out the epithelium are so deep that we have

here, as in Anguilla, distinct ducts formed between the folds

which contain on occasion large quantities of floccular matter

evidently derived from the cells. The thickened epithelium

adjacent to the rete mirabile is formed by the great ex-

tension of the folds just mentioned, the cells of which often

join at intervals, 7. e. the folds run together so that the at-first-

separate ducts form anastomosing channels before opening into

the bladder. The cells forming this epithelium are faintly

granular and almost cubical in form, and here and there contain

gas bubbles (not shown in the figure) and intracellular capil-

laries, but since I have examined these structures in greater

detail in other types, I shall not refer to them any further at

present. I shall also, and for the same reason, merely mention

here the conspicuous opacity of the cytoplasm of the gland cells

which comes into immediate contact with a capillary (fig. 19).

One other fact which is very well shown in my slides of Syn-

gnathus remains to be mentioned, and that is the disintegration

of the red blood corpuscles contained in the capillaries of the

rete mirabile and the gas gland first into spherical globules and

ultimately into granular matter. This granular matter resulting

from erythrocytolysis is, of course, only found in the blood-stream

and is to be carefully distinguished from the granular matter

above described as being found in the bladder lumen and gland

—_

GAS GLANDS OF SOME TELMOSTEAN FISHES. 195

ducts just external to the cells of the gas gland. Figure 21

(Pl. ITD.) shows several stages of this disintegration of the

erythrocytes, the significance of which I shall discuss later.

Figure 20 represents a highly-magnified transverse section

through the fine arterial and venous capillaries composing the

mass of the rete mirabile. The two kinds of capillaries are

quite indistinguishable by mere inspection. The capillaries are

separated from each other by a small amount of connective tissue.

In the very young fish, 7. ¢. before the capillaries have subdivided

to the extent found in the adult, the arterial capillaries are dis-

tinguishable from the venous, the former being thicker-walled and

somewhat smaller in calibre (see also Bykowski & Nusbaum, 28).

Thus the chief features of the second or Syngnathus type otf

“red body” are that the glandular epithelium is arranged in

simple folds, that it is restricted in area, not lining the whole of

the bladder cavity, and that the rete mirabile is contiguous with

the gas gland, although a small amount of reunion of the capillaries

of the rete may occur before these supply the epithelium.

Other Hxamples of the Syngnathus Z'ype of “ Red Body.”

GOBIUS NIGER.

As another example of the second or Syngnathus type of

bladder vasculo-glandular apparatus I shall now describe that

of Gobius niger caught at Naples*.. The “red body ” of Gobius

is a small oval red patch situated just anterior to the centre of the

oval outline of the bladder (Pl. IIT. fig. 22). It consists of a single

rete mirabile which supplies a small area of glandular epithelium.

As seen in a series of transverse sections, the artery and vein,

which break up to form the rete, at first lie owtside the several

layers of the bladder wall (Pl. IV. fig. 23) together with large

masses of pancreas. The artery and vein subdivide as usual to

form a cluster of small arteries and veins, which in Gobius, as in

Syngnathus, are intermingled with masses of pancreas (fig. 24),

1t will also be noticed that this cluster of arteries and veins is at

this stage in process of breaking through the thin dense outermost

layer of the bladder wall, so as to lie within this latter, large

Inasses of pancreas being included with the cluster of vessels,

Still more posteriorly when the arteries and veins have undergone

subdivision to a greater extent, the whole mass of vessels together

with the included portions of pancreas have become alinost

entirely encluded within the bladder wall, small portions of pan-

creas only being left outside (fig. 25). Finally, when the rete

mirabile is completely formed it is wholly included within the sub-

stance of the bladder wall, as in previous genera (fig. 26). The

rete mirabile supplies the adjacent area of pellesnidbeilors epithelium

* Gobius paganellus, obtained from Plymouth, possesses a star-shaped ‘‘ red body ”

resembling that of Peristeéhus in construction (Pl. V. fig. 37, and text-fig. 57),

though less detinite in outline. Gobdius minutus, also obtained from Plymouth, has

a similar star-shaped “red body,” but the gas gland epithelium in some specimens

is hardly folded at all, merely consisting of a single layer os cells enclosing large

capillaries ; in others it 1s folded as in Gobius niyer (Pl. LV. fig. 27).

196 . DR, W. N. F. WOODLAND ON THE

in the usual way, the epithelium lining the rest of the bladder

remaining squamous in character. It is, however, noticeable

that in the rete of Gobdiws the fine capillaries do not unite

together at all to form larger vessels before supplying the

glandular epithelium (see fig. 27). In Syngnathus it has been.

mentioned that the reunien of the fine capillaries is very slight,

but here it is quite absent.

The glandular epithelium is of the same type as that found in

Syngnathus, the cells only, as may be seen from figure 27, being

somewhat more columnar in form. The epithelial cells of Gobwus

are relatively large, and, as just stated, columnar in form and in

my preparations faintly granular. They are folded in the simple

manner already described for Syngnathus and Anguilla, large

capillaries being situated in the interior of the folds*. In figure 27

are represented three epithelial folds of the gas gland, underlying

which ave a few of the multitudimous fine capillaries composing

the rete mirabile. Even in this figure it can be observed that

several of the cells contain spherical clear spaces and that two

small capillaries have encroached on the space previously occupied

by the cytoplasm of two cells of the gas gland. ‘These intra-

cellular bubbles and capillaries and the intracellular lumina, which

1 shall also shortly mention, are cytological features which have,

within the last year or so, been studied im great detail by Jaeger

(44-47), Bykowski & Nusbaum (24), and Reis & Nusbaum (54,

55, 62-68), and they are of considerable interest from several

standpoints. I shall discuss their significance later; for the

present I may remark that I think I am justified in redescribing

these structures, especially in view of the controversy concerning

the mode of gas-production considered in Part Il. That the fine

capillaries of the rete mirabile often find their way deep into the

substance of the glandular cells, as well as between them, is proved

in Gobiws by a very moderate amount of inspection under a high

power of the microscope (Pl. IV. figs. 28, 29). It is also noticeable

that the protoplasm of that border of the glandular cells in contact

with the capillary develops a dark and often striped appearance,

which Nusbaum & Reis suggest is due to diffusion streams of

nutritive matter passing from the blood into the gland-cells, but

which I believe is intimately concerned with the production of

the gas contained in the bladder (see Part II.). Concerning the

presence of vacuolar spaces in the cytoplasm of the cells of the

gas gland, nobody can doubt their existence who has examined

well-preserved preparations of gas glands which are in an active

condition. ‘These vacuolar spaces are with good reason assumed

to represent bubbles of gas being generated within the cell-

substance (see Part Il. and Appendix B), but it is evident that only

on occasion will the gas gland be fixed and preserved when active ;

in the majority of cases, preparations ot gas glands will not

exhibit these intracellular gas bubbles, simply because the gas

%* Reis (64) states that the folds branch and anastomose in Gobius; in all my

preparations of Gobius niger, Gobius paganellus and Gobius minutus (also in

Gobius capito—Coggi, 25) the folds are quite simple, as described in the text.

GAS GLANDS OF SOME TELEOSTEAN FISUES. 197

gland is normally quiescent *. To deny the existence of these

eytoplasmie bubbles simply because certain microscopic pre-

parations do not show them is illogical, to say the least, since

intracellular gas bubbles, unlike intracellular capillaries, are not

permanent but transitory structures. Of over fifty series of

preparations which I have made of numerous types of gas gland,

not more than seven or eight show these intracellular gas bubbles

in an unmistakable manner. In the type of gas gland under

consideration, one only of my six or seven series of preparations

of the gas glands of different specimens of Gobiws niger and

G. paganellus exhibits a few gas bubbles (fig. 27). I have

seen these bubbles best in a preparation of Gobiws minutus (Pl. V.

fig. 35), in which the gas gland cells were fixed in the active con-

dition. As shown in figure 35, the majority of the gas-producing

cells possess large vacuolar spaces in their cytoplasm usually present

in the vicinity of the nucleus; indeed, the nucleus is often so

adpressed as to assume a crescentic form. These vacuoles, which,

as just mentioned, are assumed to represent the moulds, so to

speak, which contained gas bubbles, just as liver- and kidney-cells

exhibit similar vacuolar spaces which contained liquid globules

(text-fig. 60, p. 225), are of various sizes, and some can be 1 seen in

the act of being ejected from the cell substance into the bladder

lumen, where they are also occasionally found in a liberated

condition (see Appendix B). The bursting of these gas bubbles

is doubtless accountable for the masses of granular matter always

found, when the gland is active, in the bladder lumen and gland

ducts just external to the glandular epithelium, this granular

matter, of course, having composed the walls of the bubbles. The

gland cells, when active, always have that portion of their

eytoplasm situated next vascular tissue very distinct from the

rest, it being, as already described, very dense and often striated

in appearance; the rest of the cytoplasm usually assumes a

“stringy” appearance and is very vacuolater. It may also be

mentioned that the nucleus is not situated in the dense cytoplasm

next the blood, in which respect gas gland cells show a marked

difference from the cells of the pancreas, é.g.

In addition to intracellular capillaries and gas bubbles there

are also occasionally to be seen intracellular lumina or ducts—

continuations of the intercellular ducts into the substance of the

gland cells. These, again, vary greatly in different types of gas

eland and in different Specimens of each type. In none of my

preparations are they very numerous, and doubtless, like the gas

bubbles, the smaller intracellular lumina are transitory structures.

I have figured one or two of these structures in connection with

the gas gland of Ophidiwm barbatuwm, a type very effectually

studied by Reis & Nusbaum (62), and they are occasionally to be

found in my-preparations of Gobius niger and Gobius paganellus

opening into the numerous finer intercellular ducts. Since these

* See Addenda (1).

+ For excellent descriptions and figures of these gas bubbles in the cytoplasm of

the gland cells, see the papers of Bykowski & Nusbaum (24) and Reis & Nusbaum

(55, 62, 63).

Proc. Zoo, Soc.—1911, No. XIV. 14

198 DR. W. N. F. WOODLAND ON THE

intracellular channels have been so thoroughly studied by the

authors just named, I shall not refer to them in any detail. I

will merely say that since, according to Jaeger (47), gas glands

are able, when active, to swell to three times their normal size,

and that the finer ducts only become visible when filled with

fluid, it is not surprising that they are only visible in very few

preparations.

One or two other histological features to be observed in serial

sections of the gas gland and associated structures of Gobius niger

and other types remain to be mentioned. In transverse sections

across the anterior end of the rete mirabile, and only at its

anterior end, where the arteries are easily distinguishable from

the veins, all the arteries possess a peculiar endothelium. This

endothelium (PI. TV. fig. 32) contains many more nucleithan usual,

and these, instead of being flattened in the usual way, are more or

less spherical, and, each being contained in a small mass of

cytoplasm, project into the vascular lumen in the manner shown

in the figure. ‘This peculiar endothelium, in the case of the

smaller arterial capillaries, results in such a thickening of the

wall (and incidentally in some of the smallest capillaries a blocking-

up of the lumen) as to cause the capillary to bear a strong

resemblance to a bile- or pancreatic duet; indeed, were it not for

the presence of blood corpuscles these small capillaries would be

almost unrecognisable as such. This type of vascular epithelium,

which must be well-known to histologists, in all probability results

from the contraction of the pulsatile arteries in forcing the blood

through the rete, the narrowed circumference of the endothelium

causing the cells both to assume a globular form and to protrude

into the lumen of the vessel.

Concerning the posterior extension of the diffuse pancreatic

acini among the arteries and veins anterior to the formation of

the rete mirabile (fig. 25, e. g.) there is little to note other than

the fact itself, which I have not seen recorded by writers on the

teleost pancreas (see list of references to pancreas literature

below). When first studying teleost ‘red bodies” I was misled

into supposing that this extraordinary extension of the pancreatic

acini through and right into the bladder wall in many teleost

genera represented a new gland specially developed in connection

with the rete mirabile. I suggested (78) that the purpese of this

supposed new gland was the abstraction from the venous blood

stream of the globules and granules resulting from the breaking-

up of the red blood corpuscles referred to above, and in my sections

(stained by the picro-indigo-carmine method described in Ap-

pendix A) the zymogen granules present in the pancreas cells

often strongly resemble erythrocyte globules which have been ab-

stracted from the blood stream (cf. text-fig. 56 and Pl. IL]. fig. 21).

Adopting the view of Jaeger (fully discussed in Part II.) that

the cells of the gas gland produce a toxin for the purpose of

breaking up the erythrocytes and so enable themselves to absorb

more easily the oxygen in the blood stream which they sub-

sequently pump into the gas bladder, I suggested that the purpose

GAS GLANDS OF SOME TELEOSTEAN FISHES. 199

of the rete mirabile—a structure essentially consisting of the

intimate juxtaposition of the veins returning the blood from the

gas gland and carrying the hypothetical toxin, with the arteries

carrying freshly-oxygenated blood to the gas gland—was to allow

the toxin in its venous capillaries to diffuse into its arterial

capillaries and so to disintegrate the oxygen-laden erythrocytes in

time for the oxygen to be available for abstraction by the gas

gland cells by the time the arterial blood reached the gas gland.

On this view the veins of the rete mirabile would be laden with

granular matter (as indeed they are—see fig. 32) which requires

to be eliminated, and I suggested that this process of elimination

was the function of these pancreatic masses which are so closely

connected with the veins at the anterior end of the rete—the kind

Text-fig. 56 (x circ. 470).

Vein surrounded by modified acini (three shown) of the pancreas

in Nerophis equorius.

of yessels and the identical position that the hypothesis would

suggest ; in other words, the hypothesis was supported by the

position of the gland, the modification of the pancreatic acini

surrounding the veins (described below), the great similarity

between the erythrocyte globules and granules in the blood, the

zymogen granules in the pancreatic cells and the granules in the

pancreatic ducts, and the @ priori necessity for the elimination of

the erythrocyte granular matter. This hypothesis, however, con-

cerning the function of what I subsequently recognized as the

panereas is obviously untenable, since the zymogen granules are

of course present in pancreatic cells not associated with the rete

mirabile veins and, as in higher Vertebrates, do not occur outside

the pancreas cells; also the posterior extension of the teleost

14*

200 DR. W N F. WOODLAND ON THE

pancreas into the bladder wall cannot be supposed to be of any

more significance than its penetration into the liver. I failed at

first to recognize this gland as the pancreas, both because of its

peculiar position inside the bladder wall and because of the

peculiar modification of the pancreatic acini in juxtaposition with

the veins, neither of which peculiarities have been described by

writers on the teleost pancreas. Another small peculiarity existing

in several teleost genera (Globius miger, e. g.) which I studied more

than others was the syncytial character of the wall of the pancreatic

duct(Pl. V.fig. 33 and PIV. fig. 34),a feature not present in most

teleosts. The modification of the pancreatic acini in connection ’

with the veins (veins from the rete mirabile as well as hepatic-

portal veins from the gut wall) just referred to consists of the cells -

of each acinus next the thin vein wall being drawn out, the cells

on the side remote from the vein being quite short, as shown in

text-fig. 56. This figure also indicates the similarity between the

zymogen granules and the erythrocyte globules, both globules and

zymogen granules being stained a bright emerald-green in my

preparations. As stated in Part IJ., I now have good reason to

believe that the erythrocyte granular matter is abstracted from

the blood in part by the liver and in part by special cell-

masses situated near the kidneys. Laguesse (4-10), Rennie

(17), and others have shown that the teleost pancreas resembles in

all essentials the pancreas of higher Vertebrates—in the characters

of the acini, ducts, zymogen granules, and the presence of centro-

acinal cells and areas of Langerhans. With respect to these last,

IT may mention incidentally that I can fully confirm the results of

Rennie (17) and there can be but little doubt now that these

areas constitute an organ physiologically and in many teleosts

anatomically distinct from the pancreatic acini. This view of

the physiological independence of the islets of Langerhans is sup-

ported by the researches of Diamare (3), Flint (3a), Opie (14, 15),

de Witt (2), Lane (11) «nd others in opposition to the view that the

islets are patches of exhausted pancreas, supported by Lewaschew

(12), Laguesse (6, 10), and Dale (1); and, as I have already re-

marked, researches on the teleost pancreas indicate quite plainly

that the former is the correct view.

SIPHONOSTOMA TYPHLE (RONDELETII JDelar.), NEROPHIS

QUORIUS.

The ‘red bodies” of these two genera are, like that of Syn-

gnathus, small red patches situated at the extreme anterior end of

the bladder and identical in their plan of construction. The only

particular in which the “red body” of Siphonostoma differs from

that of Syngnathus is that in the former the cells of the gas gland

are much larger and the intervening ducts much narrower ;

indeed, in the posterior portion of the gland the lumina are so

narrow as not to be seen very easily. The large capillaries are

also more circular in transverse section in Siphonostoma than in

Syngnathus. Gas bubbles and intracellular capillaries are nm my

preparations to be found occasionally in the substance of the

GAS GLANDS OF SOME TELEOSTEAN FISHES. 201

epithelial cells, also the cytoplasm in contact with the capillaries

is darkened in the usual way. A small amount of granular

material is present in the bladder lumen and gland ducts just

external to the cells. I must also mention a peculiar class of

corpuscles found in the blood of my specimen of Siphonostoma

typhle. These peculiar corpuscles (PI. V. fig..36) possess very dense

cytoplasm, are about half the size of the red corpuscles, and

contain large nuclei. They are fairly numerous, forming roughly

about 1 per cent. of the total number of corpuscles. They doubt-

less represent white corpuscles, though in my preparations they

to some extent resemble small rounded ganglion cells; indeed,

they bear quite a strong resemblance to the large dense ganglion

cells so often found between the pancreatic acini, and are in con-

sequence very conspicuous. | have not observed such conspicuous

white corpuscles in the blood of any other of my preparations.

In WNerophis the gas gland epithelium is folded like that of

Gobius niger and possesses the saine features.

GASTEROSTEUS SPINACHIA.

In this Stickleback the only feature in which the ‘ red body ”

differs from those of the last four genera described is the splitting-

up of the rete mirabile into three or four bunches supplying the

gas gland epithelium at different levels; instead of the initial

artery and vein wholly dividing up to form asingle rete mirabile,

an artery and a vein are given off from this to form a rete

mirabile in connection with the mest anterior region of the gland

epithelium, another pair to form a second rete mirabile in con-

nection with the middle region of the gland epithelium, and,

finally, the initial artery and vein themselves form a third rete

mirabile to supply the rest of the gas gland. This condition

in Gasterosieus is intermediate between that described for

Syngnathus and that about to be described for Peristethus. In

all other respects the ‘‘red body” of Gasterostews resembles the

Syngnathus type.

PERISTETHUS CATAPHRACTUS (Penis J) TEDION CATAPHRACTUM

Havinys

The “red body” of Peristethus is star-shaped (Pl. V. fig. 37),

owing to the fact that the artery and vein which supply the glan-

dular epithelium, when arriving at the centre of the star-shaped

“ved body,” break up each into ten vessels. The score of vessels

thus formed sort themselves out into pairs of vessels, each pair

consisting of an artery and a vein. These pairs radiate out from

the point of subdivision of the original artery and vein like the

spokes of a wheel and give rise to ten elongated retia mirabilia in

the manner shown in text-fig. 57. Hach rete mirabile supplies a

vadially-disposed tract of glandular epithelium developed as usual

from the innermost cellular layer of the bladder wall. The capil-

laries of each rete mirabile coalesce to a certain extent to form

larger vessels before supplying the gas gland (not indicated in

202 DR. W. N. F. WOODLAND ON THE

text-fig. 57), but not to the same extent asin Syngnathus. Figure

38 (Pl. V.) represents a section (A—B in text-fig. 57) across the

anterior end of the star-shaped “red body.”

The epithelium of the gas gland is folded as in Gobius niger,

but, as in Syngnathus, it becomes considerably thickened by the

elongation and fusion of the folds in the vicinity of the rete

mirabile. The cellsare rather small and short-columnar in form,

and, in the particular specimen which I examined, contained

numerous gas bubbles (figs. 39, 40, 41); in other words, the gland

was here in a highly active condition when preserved.

Text-fig. 57.

The construction of the “red body ” of Peristethus cataphractus.

E., epithelium of gas gland; R.M., rete mirabile. Fig. 38 (Pl. V.) represents a

section taken across A-B.

The most remarkable feature about the “ved body” of Peri-

stethus is, of course, the radial disposition of the divisions of the rete

mirabile and the glandular epithelium ; in every other respect it

appertains to the Syngnathus type as above defined.

TRIGLA HIRUNDO (CoRAX Bp.).

The bladder of Trigla hirwndo, mt observed from the ventral

GAS GLANDS OF SOME TELEOSTEAN FISILES, 203

aspect, appears as an elongated sae, the ventral wall of which is

very muscular in the median line posteriovly. At the sides of this

muscular-posterior median portion of the bladder wall there are

to be seen two red streaks, which are the lateral divisions of the

“red body” (PI. VI. fig. 42), In a series of transverse sections it

can be seen that the cavity of the bladder has a peculiar conform-

ation* : anteriorly the bladder cavity consists of two laterally-placed

channels, which end blindly anteriorly and proceed posteriorly

over a distance equal to about one-third the length of the bladder

when each divides into two ; these four channels thus formed then

extend to about midway in the length of the bladder, when the

two inner channels coalesce to form a wide median chamber,

which, with the two external channels, extends to the posterior ex-

tremity of the bladder (PIL. V. fig. 43). It will be observed in figure

43 that two great median bands of muscle (M.B.) are developed

in the ventral wall of the posterior haif of the bladder, and this

explains why it is that only the lateral “red bodies” are seen

when the bladder is viewed from the ventral aspect (fig. 43).

Also shown in this figure are the two laterally-placed strands (5)

of nerve-fibres developed in connection with the two muscle-

bands.

The epithelium of the gas gland is folded in the manner cha-

racteristic of this type of “red body,” but in Zrigla it is more

folded than in previous examples (PI. VI. fig. 44), the primary

folds bearing small lateral folds. In some places the depth of the

folding is increased as in Syngnathus. The cells are of moderate

size and columnar in form. ‘he rete mirabile is divided up into

capillary tufts, 7. e. there are numerous small retia mirabilia which

supply the epithelium at intervals in its course ; in other words,

pairs of vessels are given off at intervals from the bladder artery

and vein, each pair consisting of an artery and a veinand forming

a rete mirabile to supply a certain area of gas gland.

TRIGLA GURNARDUS.

Concerning the gas gland epithelium of this species, see p. 207,

below.

SMARIS MAURII, SMARIS VULGARIS,

The “red body ” of Smaris vulgaris has the macroscopic appear-

ance shown in figure 45 (PI. VI.). Figure 46 (PI. V.) represents

diagrammatically a transverse section across the anterior end of

the “red body,” the primary artery and vein entering at the

centre of the oval mass.

Concerning the character of the gas gland epithelium of these

two species (Pl. VI. fig. 47), see p. 207, below.

Previously described “ Red Bodies” of the Syngnathus Type.

The glandular epithelia of the “red bodies” of Gobius capito,

Trigla gurnardus (see my description below), and Gasterosteus

* See description in Milne-Edwards (49).

204 DR. W. N. F. WOODLAND ON THE

aculeatus have been described and figured by Coggi (25); Gaster-

osteus aculeatus has also been described and figured by Deineka

(29); Blennius, Gobius, Syngnathus, and other types have been

briefly described by Reis (64). All these ‘“‘ red bodies” evidently

belong to the Syngnathus type as defined above.

A Third Type of Bladder Vasculo-glandular Apparatus,

as exemplified in Ophidium barbatum L.

The “ved body” of Ophidiwm barbatum has received a con-

siderable amount of attention during the last few years, especially

from the cytological standpoint (Bykowski & Nusbaum, 24 and

Reis & Nusbaum. 62, 63), since the huge cells composing the gas

gland offer exceptional facilities for enquiry in this direction.

But, as in some other instances, good figures of the gross anatomy

do not exist, so far as I am aware, and hence I shall make no

apology for recapitulating already-known facts in a manner that

shall be intelligible to the reader who has not worked at the

subject.

The “red body? of Ophidium barbatum is situated anteriorly

in the ventral wall of the bladder and has the curious appearance

shown in figure 48(P]. VI.). The fan-shaped deep red portion is, of

course, the rete mirabile, which is here an undivided body. and it is

formed by an artery and vein which enter at the pointed posterior

apex. ‘The pink margin surrounding the anterior border of the rete

is, obviously, the gas gland, A transverse section running mid-

way through the “ red body ” exhibits the parts diagrammatically

indicated in figure 49 (the outer layers of the bladder have been

stripped off). In the centre is the rete mirabile and at the sides

are the enormously thickened glandular portions of the (elsewhere)

flattened lining epithelium. As in some former instances, there is

no reunion of the capillaries of the rete mirabile before supplying

the glandular epithelium : they directly supply the gas gland.

Now in this type of ‘red body ” the gas gland does not consist

of a single layer of cells which has become folded, but of a many-

layered mass of cells, the arrangement of which bear's no resem-

blance to the folded condition of the single-layered epithelium of

the ‘“red-bodies” already described (see figs. 54, 58, 63 for

examples of the massive type of gland), and, judging from the

development of this massive type of gas gland epithelium as seen

in Atherina, it never does, at any stage of its development, bear a

resemblance to the folded type of gland. The many-layered or

massive kind of gas gland epithelium, as seen in Ophidium,

Atherina, and other genera to be mentioned, is, then, a distinct

type; the question raised by Reis in a recent paper (64) as to

whether there exist forms of bladder epithelium. transitional be-

tween these two types is one which I shall discuss when describing

Atherina. The faintly-granular cells composing this many-layered

gas gland of Ophidiwm are of quite moderate dimensions in the

vicinity of the rete mirabile, but become larger the further they

are removed, and the cells at the remote edge of the gland are

remarkable for their large size, In this type of gland there exist

GAS GLANDS OF SOME TELEOCSTEAN FISHES. 205

no large ducts comparable with those we have seen in the glands

consisting of folded epithelium ; on the contrary, the only ducts

visible are narrow slit-like channels occasionally to be detected in

their course between the cells. These narrow channels vary in

width greatly according to the state of activity of the gland; they

are necessarily numerous (though by no means always to be

observed) in order to provide means of exit to the surface for all

the numerous cells concerned, and they open on the gland surface

inside the bladder lumen by small pores. Also penetrating the .

mass of the gland are numerous fine capillaries supplying the

individual cells with nutrimert and oxygen on the sides remote

from those facing the ducts just mentioned. All the intracellular

structures previously mentioned as occurring in connection with

cells of the gas gland are here shown in a very obvious manner.

Figure 50 (Pl. VI.) shows intracellular lumina, one in longitudinal

section. As to the significance of these intracellular passages found

in gas gland cells, it is not necessary to regard them all as possessing

purely a nutritive utility as Nusbaum & Reis have supposed ; it

seems more probable, judging from their appearance in such cases

as that illustrated in figure 50, that some at least are concerned

with the passage of the gas bubble from the interior of the cell

to the exterior; they are very rare in their occurrence in my

preparations and in transverse section closely resemble bubbles,

especially if near the nucleus. The whole mass of cells composing

the gas gland is therefore penetrated by capillaries and gland

lumina. Ihave not observed a pancreas in the region of the

rete in Ophidium.

Other Examples of the Third or Ophidium Type of ‘ Red Body.”

Box sBoors L.

The ‘‘red body” of Box loops has the curious macroscopic

appearance shown in figure 51 (Pl. VI.). Figure 52 (Pl. VIT.)isa

diagram of a transverse section through the middle of the “red

body,” and the appearance of two concentric loops is thus seen to be

due to the disposition of the rete mirabile. ‘he primary artery

and vein enter the “‘red body” posteriorly and break up into

smaller arteries and veins (intermingled with pancreatic acini) to

form the rete mirabileas usual. The glandular epithelium consists,

as in Ophidiwm, of a mass of cells interpenetrated by capillaries

and lumina, but here the cells are all approximately equal in size

and the gas gland is more than a dozen cells thick.

ATHERINA HEPSETUS L,

In macroscopic appearance the “red body” of Atherina hepsetus

is diffuse in form (Pl. VI. fig. 53) and consists of tufts of retia

mirabilia borne laterally on a median pair of vessels (artery and

vein) with corresponding tracts of glandular epithelium. The glan-

dular epithelium (Pl. VII. fig. 54) is very similar to that of Lox

boops ; in other words, it consists in its thickest parts of a mass of

small cells, seven or eight cells in thickness, with inter- and

206 DR. W. N. F. WOODLAND ON THE

oceasional intra-cellular duets and capillaries. I have not observed

a pancreas in the region of the rete. The cell-outlines of the

gas gland are faint and a small amount of granular material is

present in the bladder lumen in my preparation.

I stated above that the massive type of gas gland is quite

distinct from the folded type in certain genera, and, indeed, these

two types of bladder epithelium are almost always easily distin-

guishable from each other in the genera which I have studied:

genera, with a few exceptions to be mentioned, either possess

the folded or the massive type of gland and not glands inter-

mediate in structure. In young specimens of dAtherina about

15 mm. in length, that portion of the bladder epithelium which

is about to o produce the gas gland is at first. composed of a single

layer of cubical or short Colter cells, and it is worthy of notice

that these cells, in order to build up the fully-formed massive

gland, all divide at right angles to the plane of the bladder wall

(Pl. VII. fig. 55), i.e. the successive planes of cell-cleavage are

pavallel with the plane of the bladder wall—the initial unilaminar

epithelium never becoming folded at any stage of development.

As the gas gland cells multiply connective tissue cells may be

observed to extend in between them in order to furnish the

connective and vascular tissues which penetrate hie mass of the

gland when fully formed. Also if figures 54 and 55 be compared,

it will be seen that the cells (especially the nuclei) of the gas

gland of the adult Atherina are noticeably smaller as compared

with those in the young fish. The fact that there is no indication

of folding of the bladder epithelium during development of the

massive type of gland is important, since Reis (64) supposes, on

the ground that there exist forms of bladder epithelium more

or less intermediate in structure between the folded and massive

types, that the latter has been derived from the former, and if

such be the case we might expect that the massive type of gland in

its development would pass through a folded developmental stage.

As we have seen, this is not the case in Atherina and presumably

not in other genera possessing typically massive glands. JI assume

this with thie more confidence since in some gas glands which,

owing to the exceedingly close apposition of the folds, at fist

sight appear to be massive, it is always possible to refer them

at once to the folded type by observing in each such case the

margins of the gland where it merges into the squamous epithe-

lium lining the greater part of the bladder cavity, since here the

folds are quite simple: such marginal folds are never present in

true massive glands.

Reis, in contending that the massive type of gland is but a

derivative of the folded type, instances as transitional forms

between the two types the gas gland epithelia of Syngnathus and

Girardinus. I have not examined Girardinus, but the gas gland

epithelium of my specimens of Syng gnathus can certainly not be

said to be transitional, since, as my aes iption above of this epithe-

lium clearly shows, the massiveness of the gland merely results from

the extreme length of the folds and them occasional anastomosis :

GAS GLANDS OF SOME TELEOSTEAN FISHES. 207

the glandular epithelium is always wnilaminar and the folds always

clearly recognizable. My preparations do not at all confirm

Reis’s statement that the tubular outgrowths are so numerous at

the base of the gland that they lose their lumina through mutual

contact, and form almost complete layers of epithelial cells. On

the other hand, the gas glands of some other genera which I have

examined certainly do appear to be transitional in structure

between the folded and massive types, viz. those of Zrigla gurnar-

dus, Smaris maurti, and Smaris vulgaris. I may say at once

that in all three species, especially Trigla gurnardus and Smaris

maurit, the gas gland shows distinct signs of being of the folded

type, and in the two species just mentioned the folds are quite

distinct and unmodified at the edges of the glands, but, on the

other hand, the surface of the gland is covered by a continuous

single layer of columnar epithelium (never present in the typical

folded gland) and throughout the mass of the gland the cells at

certain points lose their unilaminar arrangement*. In Z'rigla

gurnardus +, and perhaps also in the other two species, the explana-

tion of the superficial cell-layer seems to be that at the edges of

the gland the unilaminar epithelium splits into two layers, the

upper remaining unfolded and forming the superficial cell-layer

and the lower becoming folded in the usual manner (text-fig. 58),

the folds, however, anastomosing to a considerable extent and, as

Text-fig. 58.

Diagram to explain the probable construction of the gas glands of

Trigla gurnardus and Smaris maurii.

already mentioned, the cells in places becoming clustered into

groups more than one layer in thickness. In Smaris maurii (Pl. V1.

fig. 47) the individual cells are larger than in Zrigla gurnardus

and the anastomosing folds more closely packed; here and there

the cells are several layers thick, but in general they are arranged

in single folded rows as in Zrigla gurnardus. In Smaris vulgaris

the epithelium has almost entirely lost its folded character, though

this is sometimes to be detected at the edges of the gland. Thus

the gas glands of these three species{ are to a certain extent

* In many massive glands the cells are often in places arranged in single rows,

and this is evidently due to the necessity (which Oppel points out : see footnote on

page 208) of each cell being in contact on one side with a duct and on the other

with a biood vessel.

+ Coggi (25) provides a figure of the epithelium of this species showing a simple

folded character; the epithelium in my specimens more resembles his figure of the

epithelium of Motella mediterranea with the addition of a superficial covering

layer of columnar cells.

~ And possibly those of Polyacanthus. (Macropodus) described by Reis &

Nusbaum (62) and Motella described by Coggi (25).

208 DR. W. N. F. WOODLAND ON THE

intermediate in character between the folded and massive types,

but we may still retain these two categories, since we have reason

to suppose that all of these three supposed transitional forms of gas

gland really belong to the folded type and that they have only

secondarily acquired features characteristic of the massive type

by the splitting-off of a superficial cell-layer covering the surtace

of the gland, by close apposition and anastomosis of the elongated

folds, and by occasional local proliferation of the cells; in other

words, until their development is known, we may provisionally

distinguish between truly or primarily massive glands which

never exhibit the folded arrangement of cells at any stage of

development, and pseudo- or secondarily-massive glands which are

secondarily derived from the folded type*. To me Reis’s sugges-

tion that all massive glands have been derived from folded glands

seems very improbable, since on this view the folded condition

should occur as a stage in their development, and we have seen

that this is not the case in at least one typical genus (Atherina).

It seems more probable that both folded and massive types have

originated independently trom the simple unilaminar condition

which must in all cases form the starting-point of development

both in ontogeny and phylogeny.

Corts JULIS tT (VULGARIS Flem.).

The ‘red body” of Corts julis has the circular shape seen in

figure 56 (Pl. VI.), and the rete mirabile supplying the gland is

noteworthy for the way in which it is broken up into small divisions

(Pl. VII. fig. 57). The glandular epithelium (fig. 58), which is

* Reis’s classification of gas gland epithelia (64) seems to me confirmatory of

this view. Her classification is as follows: (1) entirely unilaminar epithelium

folded mto simple tubular outpushings; (2) the epithelial tolds are much folded

and branched [ Reis has made the mistake of including in this group thé gas gland

epithelium ot Corvina; in my preparatious of Corvina nigra the epithelium is of the

most typical massive type}; (8) the epithelium isin parts folded and in parts

multilaminar [I cannot agree to the inclusion of Syngnathus in this category. I

have not examined Reis’s other evample—Hippocampus] ; (4) the compact or massive

glands of Sargus, Charar, and other genera. Reis regards these four types as a

developmental series—a mistake I have corrected in the text. Reis also remarks

that only in the first three types are the ducts obviously the spaces between the

epithelial folds or tubular outgrowths ; in the fourth there exist no such intertubular

ducts, intercellular crack-like channels only being present. This last statement

seems to me to be confirmatory of my contention.

Reis makes another statement which, as Oppel (58) points out, contravenes all

probability. This statement is that whereas in the folded type of gland the

individual gland cell only presents one surface towards the duct lumen (the others

being contiguous with other cells and with a blood-vessel), in the massive gland the

individual gland cell presents all its surfaces towards the intercellular lumina pene-

trating the mass of the gland, z.e. the intercellular lumina, according to Reis,

surround each cell. Reis suggests that this supposed feature is for the purpose of

facilitating the exit of the gas through the gland into the bladder. As Oppel

justly remarks, how can acell thus shut off from all vascular tissue obtain nourish-

ment? So far as my observations go, there is no reason to suppese that a single cell

of the massive giand is not in contact with a capillary. I cannot, however, agree

with Oppel’s groundless suggestion that tubular and massive glands possess different

functions, tubular glands being secretory and massive absorptive. If this be so, from

whence do fish with massive glands obtain their bladder oxygen ?

+ Mr. Tate Regan intorms me that, according to Holt, “ Corvis giofrcdi” is merely

the female form of C. jalis.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 209

about twelve cells deep, is an example of the massive type

possessing a squamous lining epithelium. The lumina consist of

small crack-like spaces visible here and there between the cells

(largely arranged in vertical rows) in transverse sections

of the gland, and they form a system of fine anastomosing

channels opening at various points on the surface into the

bladder lumen. Gas bubbles were occasionally to be found in

the gland cells. I did not observe any intracellular capil-

laries, but they doubtless oceur; intercellular capillaries, on the

other hand, are extremely plentiful and occur largely near the

extreme edge of the gland underlying the layer of flattened

cells. The glandular epithelium of Coris julis differs from

preceding examples of the massive type in that the cells lying

next the bladder cavity form the squamous layer, or in places

layers, Just mentioned.

It was doubtless because Corning (28) chiefly examined “red

bodies” of this extreme massive type, as e. gy. that of the Pike,

that he concluded that the bladder glands possess no lumina, and,

indeed, were it not for our knowledge that most bladder glands

possess lumina and that these vary greatly in volume according to

the state of activity of the gland, we should probably come to the

conclusion if studying exclusively the same material.

CorRvVINA NIGRA Cuv., SARGUS RONDELETIY C. V., BaLisrEs

cAPRiscus L., CARANX TRACHURUS and SERRANUS CABRILLA L.

The “red bodies” of these genera all belong to the massive

epithelium type, although they vary in appearance macroscopically

(Pls. VIT. & VIIT. figs. 59-61) in correspondence with the varying

configuration of the rete mirabile. The “red body” of Balistes

alone is situated near the posterior end of the ventral wall of the

bladder. In all cases the glandular epithelium has the usual

structure—a thick mass of cells, penetrated by large or small

intercellular and intracellular lumina and capillaries.

ZEUS FABER L.

The “red body” of Zeus faber is of the curious form depicted in

figure 62 (Pl. VIIT.),and occupies the anterior half of the ventral

surface of the bladder wall. It has already been figured by Vincent

& Barnes (75). The “ red body ” consists of two lateral divisions,

each division being shaped like a C divided into three parts, and

its concavity facing that of its companion. The primary artery

and vein enter at about the centre of the “red body” and supply

branches to the retia mirabilia lying on the inner sides of the gas

gland thickenings. Anteriorly to the “red body” a thick median

longitudinal mnuscle-band is developed in the ventral bladder wall.

The glandular epithelium of the John Dory is of the ordinary

massive type, though more folded in appearance than is usual.

It is significant that Vincent & Barnes, who wished to reduce

all “red body” glandular epithelia to the folded type, remark

210 DR. W. N. F. WOODLAND ON THE

that ‘we have not been able to obtain sections which show the

direct transition to the tubular glands, but there is no doubt that

the epithelium takes on fairly suddenly a markedly glandular

aspect.”

A conspicuous feature in the gas gland of Zeus is the presence,

among cells of the ordinary size, of giant cells and nuclei six to

ten times as large (Pl. IX. fig. 65). Similar giant cells also oceur in

Ophidium barbatum, Pierasfer acus, Peristethus cataphractus,

alistes capriscus, Trigla gurnardus, and, in some instances, in

Gobius paganellus and in other genera. The only suggestion

concerning the origin of these giant cells with which I am

acquainted is that implied by Deineka (29), who states that the

similar giant cells in Perea give rise by amitotic division of the

icleus to groups of cells of the usual size—the implication being

that the giant cells are primary products of development and cells

of normal size secondary products. My observations have led me

to a somewhat different conclusion. In the first place, in many

gas glands, containing giant cells (Zeus, e. g.), these giant cells

usually lie more or less remote from the region where the blood-

vessels enter the gland—giant cells are nearly always found

towards the per ipher y or edge of the gland epithelium ; secondly,

there is every gradation in size from the smallest to the giant

gas gland cells, the smallest always being situated in those

ortions of the gland next the large blood vessels, 7, e. at: the bases

of the folds of folded glands and at the “ hub” or point of entry of

the blood-vessels in massive glands (see figs. 46, 49, 52, on Plates V.,

VL., VII. e.g.) in addition to other regions, the cells of intermediate

size like the smallest cells being found in most regions of the gland

and the giant cells, as already mentioned, towards the peripheral

portions of the gland * ; thirdly, among the smallest cells mitotic

figures are quite common in many of my preparations, but among

the intermediate and giant cells it is rare to find mitosis, though

I have seen several cases of it in intermediate cells and once or

twice even in cells approaching giant size; finally, J can confirm

Deineka in his statement that the great majority of giant cells and

many intermediate cells divide amitotically. Figure | 63 (Pl. VIIT.)

shows the appearance of the epithelium at the base of the gland of

Zeus, where the capillaries of the rete enter. It will be observed

that in this genus many of the basal portions of the massive

epithelium are syncytial in character, no cell-outlines being distin-

guishable, and that occasionally mitotic figures are present among

the nuclei of these syncytia. As we proceed from these syneytial

masses towards the periphery of the gland, cell-outlines soon become

more and more distinct and larger in size, but, as we might

anticipate, though the giant cells are usually found towards the

periphery of the gland, yet giant nuclei and nuclei of a size

intermediate between these and the smallest nuclei are to be found

* Reis remarks that in Sargus (in which there are no giant cells) the cells are’

largest next the blood-vessels anc! decrease in size towards the periphery, where they

are squamous in character. This is not the case in my preparations of Sates

rondeletii, where the cells are practically uniform in size at all depths of the gland,

except per ‘haps at the very edge, where they are squamous.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 211

in the syneytial masses at the base of the gland. In cells of

intermediate size and in giant cells amitosis occurs, as. I have

already statel. Figure 64 illustrates several phases of direct

division, but L must also mention that the huge nuclei of many

giant cells, like those of myeloplaxes of bone-marrow, appear

sometimes to divide into several—three, four, or more—nuclear

lumps at a single division, though I do not remember to have

seen cells containing more here two nuclei in the resting condition.

These various facts “considered collectively seem to suggest that in

cases like Zeus the initial unilaminar epithelium of the bladder

wall must contain nuclei of various sizes and that the nuclei of

each size, associated with cytoplasm, multiply in order to produce

the massive gland of the adult fish. As the nuclei become more

numerous they migrate from the centre of proliferation, the small

nuclei occupying cytoplasm which, owing to the size of the nuclei,

becomes subdivided up into small areas, the larger nuclei, fewer

in number, also occupying cytoplasm which, owing to the superior

size of the nuclei and their slow rate of division, remains to a

large extent unsubdivided, so forming the giant cells round the

giant nuclei. There is no evidence in my preparations of giant

cells splitting up into groups of small cells. The small cells

multiply by mitotic division, and amitosis is only found in the

ease of the larger nuclei—those of the giant cells and larger

intermediate cells. As to the origin of the giant nuclei I can

give no information. It is stated that the cells of bone-marrow

and some glands divide mitotically after undergoing amitosis, so

that there is no theoretical objection to supposing that the small

syneytial nuclei have been produced, as Deineka supposes, by

amitotie subdivision of giant nuclei, but I have never seen any

appearance of this taking place and I very much doubt its

occurrence: nearly all the : appearances of amitosis which I have

observed have been in connection with the nuclei of giant cells,

i.e. far away from the syncytia. As to the reason fen amitosis

occurring in connection with giant nuclei, it is only possible to

veadvance the old suggestion that it isa pr elude to degeneration.

The gas gland is-in Zeus and presumably other types “constantly

being regenerated at its base, and therefore there is ground for

supposing that cells at the periphery of the gland become used up

and degeneration is thenatural result of this. But this supposed

degeneration of the cells has nothing to do with the violent

decomposition of cell-substance assumed to occur by Nusbaum &

Reis in connection with the production of gas—a phenomenon for

the existence of which I can find no evidence and in which I

quite disbelieve.

GADUS MORRHUA.

The “red body” of the Cod* is known to every student of

comparative anatomy as a large red mass situated anteriorly in the

bladder. As Vincent & Barnes (75) say, the free surface of the

“red gland” shows “many small clubbed processes, packed

* T used common “Codling” caught off Deal.

2) DR. W. N. F. WOODLAND ON THE

tightly together, each consisting of a red mass of tissue, sur-

mounted by a cap of yellowish mucilaginous-looking material.”

Each club-shaped process ‘“ consists essentially of two parts :—

(1) a mass of capillary blood-vessels, lying side by side, conveying

blood te and taking blood from the second parts; (2) a glandular

portion.” In figure 66 (Pl. VIII.) is represented diagrammatically

a transverse section of the gland of the ‘‘ Codling,” which shows

the “clubbed processes” well in section, also one “cap of

mucilaginous-looking” connective tissue. Vincent & Barnes go on

to say that “when the capillaries arrive at a point about -2 mm.

from the free surface of the gland, they interdigitate with

involutions of the glandular epithelium, which dips down between

them and forms a covering tor them, thus constituting tubular

glands -2 mm. in length.” In other words, Vincent & Barnes

state that the glandular epithelium of Gedus is that which I have

deseribed above for the Hel and Syngnathus, and they provide a

diagram of what they suppose to be the folded arrangement of the

epithelium. Iam quite willing to admit that the small columnar

cells which compose the glandular epithelium are in general

arranged round strands of vascular connective tissue, and that

occasionally the intercellular lumen is so extensive and so disposed

as to give the glandular tissue an almost folded appearance, but

there is no real ground for comparing the massive structure of the

glandular epithelinm of the “ Codling” to the folded epithelium

of Syngnathus, The gas gland of the “ Codling” 1s of the massive

type, in which the intercellular lumina are very abundant and

occasionally very large. The capillaries are very small and run

in the numerons thick strands of connective tissue. As Vincent

& Barnes mention, caps of connective tissue, continuous with

the connective tissue penetrating the glandular epithelium, are

often present. I have not examined them in detail.

CEPOLA RUBESCENS L.

The “red body” of Cepola rubescens 1s situated anteriorly in

the ventral wall of the bladder and has somewhat the shape of a bell

(Pl. VIII. fig. 67), the handle being formed by the rete mirabile.

The artery and vein enter the “red body” anteriorly at the top of

the bell-handle, and there give rise to the rete mirabile in the usual

way. The rete mirabile gradually diminishes as it supplies the

mass of the gas gland, which first appears, in the series of trans-

verse sections, on the right side of the “red body” (text-fig. 59,

G.H.). The peculiar feature about the “red body” of Cepola

is the curious folding of the glandular mass anteriorly. The gas

gland extends a considerable way posteriorly, as seen in figure 67,

and the best way to comprehend the folding anteriorly—the

folding, that is to say, of the whole mass of the gas gland, not

the folding of the epithelium hitherto described —is to observe the

diagrams of the sections contained in text-figure 59 in the reverse

order, 7. ¢. observe H first and A last. Diagram H represents the

glandular epithelium (the dotted area G.H.) as a mass of cells

developed from the general squamous epithelium (/.#.) lining the

GAS GLANDS OF SOME TELEOSTEAN FISHES. Dales

bladder cavity (B.C.) in the median line of the ventral wall of the

bladder. In diagram G it is to be observed that the squamous

epithelium on each side of the glandular mass has become folded

(f.F.#.) and projects into the bladder cavity towards the median

line. In diagram F these folds have nearly met in the middle

line and lie dorsal to the gas gland in the bladder cavity ; also it

will be observed that on the left side the glandular mass itself has

become completely reflected towards the median line (/.L.G.G.),

and lies in close apposition with the rest of the glandular mass.

Still more anteriorly, the two lateral folds of squamous epithelium

have, as shown in diagram EK, united in the median line, so that

Text-fig. 59.

Diagrammatic transverse sections through the anterior end of fig. 67 (PJ. VIII.),

where the rete mirabile supplies the epithelium of the gas gland. The diagram

is fully explained in the text.

two separate epithelia now lie in the bladder cavity above the gas

gland, the upper of which (/./.Z.) is continuous with the squamous

epithelium lining the rest of the bladder wall, and the lower

(O.F.Z.) is continuous with the two sides (the left reflected as

just described) of the glandular mass, which latter therefore now

forms the thickened ventral wall of a tube. Diagram D shows

that the dorsal thin wall of the tube just mentioned is being

replaced by extensions of the glandular mass, this replacement of

the thin epithelium being chiefly due to the increase in size of the

Proc. Zoot. Soc.—1911, No. XV. 15

214 DR. W. N. F. WOODLAND ON THE

reflected left side of the glandular mass. In diagram C the

reflected left side of the gas gland has increased so as to extend

over the opposite or right side of the gland, a narrow crack

(continuous posteriorly with the bladder cavity) only being left

between the dorsal and ventral halves of the gas gland... In

diagrams B and A, owing to the coalescence of the glandular

masses of the dorsal and ventral portions of the gas gland on the

right side (7. e. the replacement of the short strip of thin epithelium

which in © closed the tube on the right side), the narrow crack

this crack narrows anteriorly until it disappears shortly before

the gas gland itself terminates.

The mass of glandular epithelium (Pl. IX. fig. 68) resembles that

of Coris julis in being (apparently) almost devoid of gland Jumina.

Small intercellular cracks are visible here and there between the

cells in my preparations, but that is all to be observed in the inactive

gland. On the other hand, the glandular mass is penetrated by

capillaries, and the large cells appear to arrange themselves round

these capillaries to some extent, giving the gland in section almost

a lobulateappearance. The cytoplasm of the cells is conspicuously

darkened where in contact with the capillaries. Now and again

large cells are found with a very large nucleus, but these are rare.

Finally, it may be mentioned that the capillaries of the rete

mirabile possess the peculiar endothelium which I have already

described in Gobius paganellus and found in most “ red bodies.”

PERGA FLUVIATILIS.

The ‘ved body” of the freshwater Perch is a diffuse organ

situated anteriorly on the ventral and lateral portions of the

bladder wall (Pl. TX. fig, 69). The retia mirabilia are seen as small

fan-shaped tufts present at the extremities of the various branchings

of the closely-associated artery and vein (both included in the

red streak seen in the bladder wall), and bordering these tufts of

rete are the areas of gas gland. ‘The gas gland is of the ordinary

massive type, but feebly developed compared with the gas glands

of most marine fish, consisting in its thickest parts of not more

than six cell-layers (see text-fig. 62, p. 240) and dwindling to one

at its edges (P]. TX. fig. 70). The giant cells described by Deineka

in young Perch are only of very moderate size in my preparations *,

indeed, not so large as those I have figured in the case of Zeus ; as

in Zeus, amitosis occurs in connection with most of the cells.

With reference to the suggestion of Deineka that these giant cells

‘subdivide into groups of the smaller cells as a consequence of the

fragmentation of the nucleus, I have already advanced reasons for

regarding this as improbable (see Zews).

Previous Descriptions of the Structure of the Ophidium

: or Third Type of ‘‘ Red Body.”

I shall here merely enumerate the chief histological descriptions

of gas glands of the massive type. Emery 1880 (33), Mierasfer ;

* My Perch were from 10-14 cm. in iength.

GAS GLANDS OF SOME TELEOSTEAN FISIIES. 215

Coggi 1886 (25), Serranus, Perca, Balistes, Corvina, Julis, Motella

and other types, and 1889 (28) Lsou lucius; Corning 1888 (28),

fisox, Perca and other types; Vincent & Barnes 1896 (75),

Zeus, Gadus and other types; de Seabra 1897 (31), soa and others;

Jaeser 1903 (44), Sciena, Lucioperca; Deineka 1904 (29), Perca,

Hsox and other types; Bykowski & Nusbaum 1904 (24),

Fierasfer, Ophidium; Reis & Nusbaum 1905, 1906 (62, 63),

Polyacanthus (Macropodus), Kierasfer, Ophidium, Charax, Perca ;

Reis 1906 (64, 65), Sargus, Charax, Pagellus, Corvina, Umbrina,

Chrysophrys and others described in a general way.

Summary of Part I. and General Conclusions respecting

bee Gas Gland Structure.

From the foregoing it is now manifest that all teleostean ‘“ 1ed

bodies,” ¢. e. the vasculo-glandular gas-producing organs situated

in the bladder wall, are composite structures consisting of two

distinct and sometimes separate parts—the glandular epithelium

or gas gland* and the rete mrabile +. Vhe mutual relationships

of these two parts are quite constant, and though the gas gland

must be described ag the essential part, yet the rete, on account

of its constant = presence and undoubtedly highly important

function, is probably of almost equal importance. The rete

mirabile bipolare geminum (Miiller, 53), 7. e. the arterial and

venous rete already described, is always formed in connection

with the artery and vein supplying the gas gland, and may either

be anatomically quite distinct from the glandular epithelium (Hel

type), the arterial and venous capillaries reuniting to a varying

extent on the gland side of the rete to form large arteries and

veins before supplying the glandular epithelium, or the rete

mirabile may be contiguous with the glandular epithelium, the

capillaries of the rete undergoing little (e. g. Syngnathus) or no

(e. g. Ophidium) reunion before supplying this.

The diffuse pancreas, when present, is often intermingled with

the arteries and veins situated at the extreme anterior end of the

rete, and so actually becomes situated inside the bladder wall ; it

was at first, on account of several peculiarities, mistaken by me

for a new gland with a special function connected with the gas

gland, but this view was quite a mistaken one, the penetration

of the pancreas into the bladder wall being of no physiological

significance.

The gross anatomical differences, often visible to the naked

eye, which are to be found in ‘‘red bodies” taken from different

fishes, are determined by the disposition and degree of subdivision

of the rete mirabile—since the distribution of the glandular

* The “ driisige siiume ” of Miiller and the “ corpo epitheliale ” of Coggi.

+ ‘The “ vaso-ganglion ” of Owen and “ organo vascolare” of Emery.

{ That is to say, using the term “gas gland” in the usual sense as equivalent to

‘oxygen gland” (see Part II.), the rete is constantly associated with it; the rete,

however, is absent in the cases of those bladders the lining-epithelia of which only,

or principally, “ secrete” nitrogen and carbon dioxide.

Tor

216 DR. W. N. F. WOODLAND ON THE

epithelium must evidently correspond with that of the rete

mirabile. It need hardly be pointed out that the gross ana-

tomical simplicity or complexity of the “red body” by no means

always corresponds with the histological simplicity or complexity

of the gland epithelium, and that this latter affords the only true

basis for a classification of “‘red bodies.” One of the most simple

(though not the most primitive) conditions of the rete mirabile

is when it is single and compact*. In this condition it may be

separate from the glandular epithelium and ovoid in shape

(Ophichthys imberbis, fig. 9, e.g.), or contiguous with the glandular

epithelium and circular in form (Godius, fig. 22, e. g.), conical

(Cepola, fig. 67, e. g.) ov fan-shaped (Ophidium, fig. 48, e. g.), &e.

In some cases the rete mirabile is divided into two, and here

these are separate from the glandular epithelium and ovoid in

form (Anguilla, text-fig. 53, Myrus, Murcena, Ophichthys serpens,

&e.). In other cases the rete mirabile is divided up into several

distinct parts, which often have a radiate arrangement: e. g. 10

Peristethus the rete is split up into some ten radiating strands

(fig. 37 and text-fig. 57) and in some species of Gobiws into seven

or more. In most cases, however, the initial artery and vein

supplying the bladder give off numerous arteries and veins in pairs

(an artery and a vein to each pair, of course) at intervals so as to

give rise either (@) to continuous sheets of rete mirabile, so to

speak, very often two in number, one on each side, as in Corvina

nigra (fig. 59), Sargus rondeletii (fig. 60), and Smaris vulgaris

(fig. 45), or (6) to sheets divided up im variable degrees, as e.g. in

Box boops, where the sheets are divided into outer and inner on

each side (figs. 51, 52), or in Zeus faber, where there are three

separate sheets—anterior, median, and posterior—on each side

(fig. 62), or (c) to numerous separate small strands or tufts of rete

mirabile, each strand supplying a small area of glandular

epithelium: thus, in Coris julis (figs. 56, 57) the rete is seen

under the microscope to be subdivided into many small strands

supplying the small area occupied by the gas gland ; in Trigla

hirundo also a similar condition exists, but here, indeed, the twigs

extend nearly over the whole interior of the bladder cavity (fig. 43)

instead of being restricted to a small area; in other cases where

the tufts are larger and more separated, these can be seen with

the naked eye, as, e. g., in <Atherina hepsetus (fig. 53) and Perea

(fig. 69) and in the well-known case of the Cod (fig. 66), in which

the subdivision of the rete mirabile reaches its maximum.

Whether the condition of the rete mirabile with which we

started, 7. e. aS a single large well-defined body, separate from the

* Reis (64) supposes that the horseshoe-shaped gland of Ophidiam, e. g. (see my

fig. 48 and compare it with figs. 59, 60, and 62), is the form of gas gland and rete

from which all others are derivable, but it seems to me that the simple oval rete of

Syngnathus in connection with an extensive area of glandular epithelium, is still

more simple; the most primitive condition of the epithelium and associated blood

vessels is, of course, that of many freshwater teleosts in which practically the whole

of the internal unilaminar lining-epithelium of the bladder constitutes the gas gland,

and the arterial and venous capillaries have not become associated to form a rete.

In general it may be said that the more d:ffuse the rete and gland the more primitive

their nature.

GAS GLANDS OF SOME TELEOSTEAN FISHES. pt

glandular epithelium (Ophichthys imberbis, e. g.), is more primitive

or more evolved than the condition which we found in the Cod,

e.g., is a question I shall not discuss; I will merely say that there

are several reasons for accepting, with de Seabra (81), the latter

view. The fact that Ophichthys and Anguilla possess a very simple

type of glandular epithelium is no objection to this view, since

the rete of Cepola and other genera possessing complex types of

epithelium is almost as simple.

The glandular epithelium, as we have seen, may line the whole

interior of the bladder (as in many Anguilliformes and fresh-

water teleosts) or be restricted to a small area of the internal

surface (Ophidiwm type). The Syngnathus type is more or less

intermediate between these two conditions.

The character of the glandular epithelium varies greatly. The

cells composing it may be arranged to form (a) a single unfolded

layer (Alyrus, e. g.); (6) a layer thrown into simple or complex

folds, so that channels are formed leading from the surface of the

glandular epithelium to the bladder lumen (Anguilla, Gobius,

Syngnathus, Trigla, Smaris); (c)a thick mass in which the deeper

cells can only communicate with the bladder lumen by a system

of large or small anastomosing channels (the intercellular lumina)

which penetrate the cell-mass and open into the bladder cavity

by small apertures situated on the internal surface of the

glandular mass (Ophidiwm, Coris, e. g.). At present neither

embryology nor comparative anatomy provides evidence to justify

the inference that this massive type of gland epithelium is in all

cases derived from the folded condition—the folds on this view

being supposed to have become elongated, laterally branched

perhaps, closely apposed and fused at intervals to produce the

cell-mass penetrated by narrow anastomosing channels. It would

sratify me to be able to agree with Reis (64) who states that she

is able to trace a complete developmental series of complications

in the structure of the gas gland epithelium, the simple unfolded

eell-layer forming the starting and the cell mass of Coris, Cepola,

or Zsow the culminating points of the series; but, as I have already

stated, the available embryological and anatomical evidence

suggests that the two complex types of folded and massive gas

gland epithelia were separately derived from the primitive layer

of glandular cells by folding (7. e. the plane of cell-cleavage is

always at right angles to the plane of the bladder wall) and by

thickening (the plane of cell-cleavage being parallel with the plane

of the bladder wall) respectively. It is true that forms of gas

gland epithelia occur apparently transitional in character between

the folded and massive types (Zrigla gurnardus, Smaris maurit,

S. vulgaris), but I have supplied reasons for regarding these as

glandular epithelia of the folded type which have become

secondarily massive in form and so differing from the truly

massive glands which never pass through a folded stage in their

development. There is no reason, however, why certain glands

should not be truly intermediate in structure, ¢. e. in which the

cells during development divide simultaneously in planes both

218 DR. W. N. F. WOODLAND ON THE

perpendicular to and parallel with the plane of the bladder wall,

but I know of no examples of such. Subsequent embryological

enquiry can alone prove which of these views is the correct one.

The minute structure of the cells of the gas gland epithelium is

doubtless constant throughout the entire series of “red bodies.”

Gas bubbles are in all cases generated in the interior of the cells

(usually in the neighbourhood of the nucleus) when the glands

are active, and these bubbles are ejected by the cells into the

gland ducts and bladder cavity where they explode, the shattered

walls of these bubbles giving rise to the masses of granular matter

already described. Vincent & Barnes analysed this granular

matter (which, as before mentioned, must be cavefully distinguished

from the disintegration products of the red blood corpuscles alone

found in the blood-vessels) of the Cod and found it to consist

principally of a nucleo-proteid; and since the cytoplasm which

forms the wall of the gas bubble is also composed for the greater

part of this substance, the result of this single analysis is con-

firmatory of the view just expressed, as is also the fact that a

large amount of this granular matter is always associated with

the presence of numerous gas bubbles. The intracellular ducts,

which, with the other cytological features of the gas gland, have

been so well figured by Bykowski & Nusbaum (24) and Reis &

Nusbaum (62) within the last few years, are doubtless also constant

features of active teleost gas glands, though I am of opinion that

some of these intracellular channels are due to the expulsion of

bubbles from the cells and not purely nutritiveas Reis & Nusbaum

suppose. The intracellular capillaries, on the other hand, are

permanent structures, and the fact that they occur so rarely as

compared with the enormous number cf these structures found in

connection with the liver cells can only be explained by the

relative inactivity of the teleost gas gland.

I have already described and figured the hemolytic disintegra-

tion of the red blood corpuscles (erythrocytolysis as it may be

called) which occurs in the blood-vessels associated with the

active gas gland of Syngnathus acus, and this same phenomenon

is to be found in connection with other of my preparations of

active gas glands and is doubtless a result of the gland’s activity.

In this process of hemolysis many of the erythrocytes become

entirely broken up, first into large globules and finally into the

granular matter which is present in such large quantity in the

capillary lumina, whilst others are distorted in shape and only

partially broken up, looking very much as if influenced by some

toxic substance, as Jeger suggests (see Part I1.). This condition

of the erythrocytes cannot be due to imperfect fixation since

great care was exercised in the fixation and preservation of these

gas glands; moreover sections through other vertebrates— Elas-

mobranchs e. g.—fixed by similar methods, exhibit the blood

vessels as practically free from granular matter (the small amount

sometimes present simply representing coagulated plasma) and

the erythrocytes as retaining their form.

Another fact which I wish to lay stress upon, especially in view

GAS GLANDS OF SOME TELEOSTEAN FISHES. 219

of the theory advanced in Part I1L., is the distribution of the

granular matter in the capillaries of the rete mirabile. Careful

examination of my sections* of Gobius niger, Gobius minutus,

Syngnathus acus, Peristethus cataphractus, Gasterosteus spinachia,

and some others, in all of which the gas glands are more or less

active, had led me to the conclusions that granular matter is

present in the venous capillaries of the rete to a far greater extent

than in the arterial, and that the relatively small amount of

granular matter present in the arterial capillaries is chiefly

situated next the distal pole of the rete—the pole next the gas

gland, The small calibre of the arteries compared with the veins

and the few series of sections of active glands, well fixed and well

stained, which I possess render it difficult to be absolutely certain

of the above conclusions, but I am as certain as the limitations of

my material will permit me to be. The bearing of these facts on

the theory already outlined on page 198 is perhaps already

apparent, but I shall reserve discussion for Part IT. (see p. 232).

As already implied, teleost gas glands aye best classified

according to the character of the glandular epithelium. We

divide gas glands into two groups :—(1) gas glands in which the

glandular epithelium is a single layer, which may either remain

unfolded or become folded in vaviable degrees; and (2) gas glands

in which the glandular epithelium is many lay ered, the mass of

cells being penetrated by fine ducts. With the exception of a few

cases (7 ae gurnardus, Smaris maurti, S. vulgaris, and some

others) in which the much-folded epithelium has secondarily

assumed a massive character, this massive type of epithelium is

probably not derivable from the folded type. Also in this second

group of gas glands the rete mirabile is always contiguous with

the glandular epithelium, reunion of the capillaries never

occurring (Ophidium, Pierasfer, Box, Atherina, Coris, Balistes,

ar gus, Cemanes Charax, Serranus, Zeus, Gace, Cepola, EHsoa,

Perca, &e.).

Group | is subdivisible into two minor groups :—

(a) The rete mirabile of the gas gland consists of either one

or two bodies distinctly separate from the glandular

epithelium, the capillaries of the rete reuniting to form

large arteries and veins before they come into con-

nection with the glandular epithelium (Anguilla,

Myrus, Ophichthys, Murcena, &e.).

(6) The rete mirabile of the gas gland is contiguous with the

glandular epithelium, although a small amount of re-

union of the capillaries occasionally oceurs (Syngnathus,

Gobius, Siphonostoma, Nerophis, Gasterosteus, Peri-

stethus, Trigla, &c.).

In concluding this account of the practical work connected with

* Observations of the rete mirabile, when sectioned longitudinally, are preferable

froin this standpoint ; the arterioles are, of course, distinguishable from the venules

by thicker walls, by small ealibr e, and by the peculiar endothelium before mentioned

220 DR. W. N. F. WOODLAND ON THE

the present paper, I wish to express my great indebtedness to the

Council of King’s College, London, for generously allotting me

the sum of thirty pounds out of the Government grant to the

College for 1907 to defray a part of the expenses incurred by me

during my visit to the Naples Zoological Station in April of that

year. I wish also to express my indebtedness to the Naples

Zoological Station Table Committee of the British Association for

the occupancy of the table at Naples during the three weeks just

referred to, and-to the staff of the Naples Zoological Station for

the large amount of trouble they took in supplying me with the

material I required in connection with this and other subjects.

Further, I wish to thank Professor A. Dendy, F.R.S., and

Professor 8. J. Hickson, F.R.S., for kind recommendations in

connection with the above-mentioned grant and occupancy of the

Naples table respectively, Professor J. P. Hill, D.Sec., and Dr. G.

C. Chubb for recent friendly assistance, Mr. C. Tate Regan, M.A.,

for kindly advising me in connection with the systematic nomen-

clature adopted in this paper (the nomenclature of Giinther’s

Catalogue), and Mr. C. Biddulph, who has considerably lightened

the burden of preparing some hundreds of slides. Finally, I am

also much indebted to the Council of the Royal Microscopical

Society for kind permission to occupy their table at the Plymouth

Marine Biological Laboratory during a week in August, 1909.

Part [1.—A FEW SUGGESTIONS CONCERNING THE PHYSIOLOGY

oF THE “ Rep BopiEs.”

It was thy original intention to include in Part II. of this paper

a résumé of our present knowledge of the physiology of the gas

bladder *, but although, with this end in view, I have become

thoroughly acquainted with the whole of the vast literature

dealing with the subject, yet other work prohibits me at present

from thus devoting the time necessary for the composition of

such a review. This being the case, I shall content myself with

providing an outline sketch of the chief views which have hitherto

been advanced to explain the mode of working of the “‘red bodies,”

in order that the reader may be in a position to estimate the

value of the few additional suggestions I have to make.

The several views which have been held concerning the origin

of the threey principal gases—oxygen, nitrogen and carbon

dioxide—contained in variable proportions in the gas-bladder

cavity, can be classified into two categories: (1) the view (first

attributed to Redi) that these gases are derived directly from the

atmosphere, and (2) the views (lineal descendants of Needham’s

secretion theory t) that these gases are derived more or less

* A yecent account of tke functions of the bladder will be found in Baglioni (18).

+ Argon is also stated to occur in the bladder.

t It should be noted that authors in discussing the production of gas employ the

term “secrete” in a very loose manner, some thereby meaning a true process of

‘secretion, such as that which occurs in the sebaceous gland (Nusbaum & Reis,

e. g.), others (Hiifner, Jaeger) merely meaning a process of pumping from the blood.

GAS GLANDS OF SOME TELEOSTEAN FISHES, Al

directly from the blood stream. The first view, abandoned by

nearly all modern authors, is, however, persistently maintained

by Thilo(70, 71, 72). Thiio’s principal contention, stated briefly,

is that the blood of a fish is not sufficient in quantity to contain

the amount of gas found in the bladder, and that even if this

were the case the circulation of the fish is so feeble that the blood

could not replenish an emptied bladder in the time experiment

proves that it can be replenished. He therefore asserts that in

all cases the bladder-gas must have been procured by the fish

directly from the atmosphere: Physostomi can always renew their

bladder-gas by rising to the surface of the water and passing air

along the ductus pneumaticus, whilst Physoclisti are apparently

under the hard necessity of absorbing a sufficient supply when

young, and therefore before the duct has degenerated, to last

them throughout life, though Thilo further maintains that in

many cases even adult Physoclisti are still able to procure fresh

supplies by passing air, either swallowed from the atmosphere or

extracted in some inexplicable manner from the water, along the

strand of tissue representing the vestigial duct. Thilo performed

experiments and, according to his statements, obtained results

which strongly support his view. He asserts, e. g. that he cut off

the vascular supply of the previously-emptied bladder in inca (a

physostome) and that in thirty hours it became refilled with gas.

Seeing that the blood-supply was absent, he argues that this fresh

supply of gas must have been obtained from the atmosphere by

passage through the pneumatic duct. I, however, cannot find

that he analysed the gas produced under these conditions, and,

since atmospheric air and secreted gas nearly always consist of

oxygen, nitrogen and carbon dioxide associated in very different

proportions in the two cases, his contention as to the source of the

gas receives very little real support from his experiments. A few

of the many obvious objections to Thilo’s hypothesis may be

stated. In the first place, the percentage composition of the

three gases contained in the bladder is, as just mentioned, quite

different from that present in the atmosphere, and in the cases

of oxygen and nitrogen these gases are often present in such

quantities as to exert a pressure many times greater than the pres-

sures they exert in air—both of which elementary facts are fatal

to Thilo’s view. Thilo’s contention that the blood of a fish with

‘its feeble circulation is incapable of providing the quantity of gas

required by the bladder of course begs the question at issue, and

he might with equal force contend that the tissues of a siliceous

sponge, for example, must be incompetent to deposit a massive

siliceous skeleton, seeing that 100,000 parts of sea-water contain

little more than one part of silica in solution and that diatoms

and other minute organisms are serious competitors even for this

small quantity (Sollas). Further, Thilo’s experimental results are

not in accordance with those obtained by Hiifner (38), Jaeger (45),

and others, and, indeed, Thilo’s own experiments failed to render

his contention even probable since the gland cells of the bladder

would not necessarily cease their activity immediately a large

22 DR. W. N. F. WOODLAND. ON THE

part of their blood-supply was cut off. It is also certain that

most physostomous fishes do not obtain their gases from the

atmosphere wd the pneumatic duct, since the pneumatic duct is ©

now known to be merely one of the two principal mechanisms

(the other being the ‘‘ oval”) employed by fish for the elimination

and not the obtainment of gas. Finally, Thilo’s theory fails to

account both tor the degeneration of the ductus pneumaticus in

Physoclisti (since, according to Thilo, it is still functional) and

for the presence of the complex ‘red bodies” which are the very

structures needed by the fish on account of the feeble blood-

circulation referred to by Thilo.

= Theories which derive the bladder gases from the blood stream

can again be grouped into two classes, viz. those which suppose

that the blood gives up its gaseous constituents more or less

directly. to the bladder, the gas passing straight from the

capillaries of the rete mirabile, or wall of the bladder, into the

bladder lumen, and those which regard the gas gland as the

special organ which extracts the gas from the blood. The first

class of opinions, to some extent associated with the name of

Moreau, is now quite out of date, though still to be found stated

in some recent text-books. The second class of opinions comprises

two quite distinct views as to the exact function of the gas gland

—two views which at the present time are held with equal tenacity

by the schools represented by Jaeger and Nusbaum & Reis

respectively. I shall first state briefly the view of Jaeger (44-48).

Jaeger, following Hiifner (38), holds that the gas gland is

primarily a pumping apparatus, that is to say, an apparatus for

pumping the gases contained in the blood into the bladder cavity.

The pressure exerted by the gases in the blood is, of course,

considerably less than that exerted by the gases in the bladder ™%,

and it is the function of the gas gland to force the gas from the

blood into the bladder lumen against this superior pressure.

Jaeger further supposes that the disintegration of a certain

percentage of red blood corpuscles is effected by the secretion on

the part of the gas gland cells of a toxin which is poured into the

blood for this purpose. The object of so breaking up the ery-

throcytes is to enable the gas gland cells, in some way not

described, to lay hold of, with greater facility, the oxygen thus

scattered in the corpuscle fragments. Indeed, Jaeger, following

“Moreau, regards the gas gland as a mechanism essentially con-

cerned with the pumping of oxygen—as an oxygen gland, in

short,—a view confirmed both by the great development of this

glandjand by the disintegration of the erythrocytes in connection

with bladders containing a large percentage of oxygen, and also

by the contrary fact that in the bladders of Cyprinoids and many

other freshwater fish which mostly contain nitrogen, the gas

glands are absent—the ordinary squamous epithelial lining here

being capable, without undergoing any special modification into a

* BH. g. Jaeger (46) states that in deep-sea fish the partial pressure of the oxygen

in the blood only amounts to about one-fifth of an atmosphere, whereas the oxygen

in the bladder may possess a partial pressure of over forty atmospheres.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 228

gland, of pumping in the nitrogen alone required. The reason for

the special development of the oxygen gland in many deep-water

fishes * is because oxygen alone among the gases present in the

bladder is capable (in virtue of its property of combining in large

quantity with the hemoglobin of the blood) of being either rapidly

produced (by the gas gland) or rapidly absorbed (by the “ oval ’ 7),

and this rapidity of production and absorption is essential in the

case of fishes which undergo considerable changes of pressure in

the bladder. Deep-water marine or freshwater fish differ from

most freshwater fish in that living in great depths of water those

possessing migratory habits in a vertical direction require ap-

paratus for adapting the volume of gas in the bladder to the violent

changes of pressure experienced; most freshwater fish, on the

other hand, live in comparatively shallow water—in inland lakes

and rivers—and thus lead placid lives, experiencing little or no

changes of pressure in the bladder, and for these fish it is evident

that there is no need for the special development of an oxygen

gland, which makes extravagant use of the gas which the fish

requires for respiration: nitrogen and carbon dioxide suffice ¢.

With reference to the spherical spaces present in the cells cf the

eas gland, Jaeger altogether denies that they represent gas

bubbles: they are merely vacuoles such as are to be found in

liver-cells, e. g. and have no connection with the production of

gas. Jaeger himself, however, describes gas bubbles as being

present in the ducts of the gas gland. Jaeger also refuses to

admit that the large amount of granular matter found outside

active glands in the bladder lumen is the product of the gas gland

cells: in short, represents the broken-down walls of exploded gas

bubbles. IJ shall discuss these features of Jaeger’s theory later.

Nusbaum & Reis (64, 55, 62-66), relying upon their extensive

investigations of the cytology of the gas gland, formulate certain

peculiar views § as to the exact mode of function of the gas gland.

Jaeger, as already stated, holds with Hiifner that the gas gland

is essentially a pumping apparatus, though he confesses entire

ignorance as to exactly how the individual gas-producing cell

* Only those deep-water fishes which migrate vertically and thus experience

differences of pressure; fish which remain at one depth, however great, obviously

have no more need for “ red bodies” than have surface fish (see p. 228).

+ Any sort of gas can, of course, be eliminated by the pneumatic duct. y

+ Mile. C. M. L. Popta in her recent paper entitled “ Etude sur la Vessie Aérienne

des Poissons”? (Ann. Sci. Nat. Zool., t. xii, 1910, pp. 1-160) comes to the conclusion

that “ la vessie aérienne des poissons se développe pour aider a la circulation de

Yoxygéne [et en général des gaz] dans le corps du poisson.” I confess that this

conclusion does not convey much meaning to me. I entirely fail to understand why

the fish should develop an organ for tle sole purpose of liberating oxygen into the

blood in the same way I suppose that the liver liberates food material, and this is the

only intelligible proposition which the author’s conclusion conveys tome. Though

the bladder undoubtedly in many cases performs this function incidentally, yet the

fact that the bladder often contains other gases (nitrogen and carbon dioxide) the

presence of which in the blood cannot be of the slightest advantage to the animal,

shows that this storage function of the bladder cannot be its primary function. I

caunot see that the author’s couclusion in any way explains the different proportions

in which the three principal gases occur in the bladder.

§ Unfortunately adopted in the last edition of Wiedersheim’s ‘ Vergleichende

Anatomie der Wirbelthiere, Siehbente Aufiage, 1909.

224 DR. W. N. F. WOODLAND ON THE

extracts the gas from the blood and forces it against great

pressure into the bladder: the supposition is, on his view, sumply

a legitimate deduction from the facts, and the details of the

process, upon which he can throw no light, are of secondary

importance. Nusbaum & Reis, on the other hand, profess

to have discovered in the cytology of the gas gland cell, some

details of the process which prove that the gas gland is not the

mere pump which Jaeger assumes it to be. ‘The view of Nusbaum

& Reis is briefly this :—Hxamuination of the cells of the gas gland

proves the existence of spherical spaces in the cytoplasm which

are not mere vacuoles as Jaeger assumed, but represent actual

gas bubbles being formed in the cell. These gas bubbles are also

to be found in the act of being ejected from the cells, and also

lying freely in the gland ducts and bladder lumen, Jaeger himself

necessarily agreeing that they are bubbles when found outside the

cells, since liquid vacuoles could not possibly so persist. Thus

much is a matter of observation, but Nusbaum & Reis contend,

in explanation of these facts, that these bubbles of gas are pro-

duced by the actual decomposition of the substance of the gas gland

cells, just as gases are produced by putrefaction, and in proof of

this cell-decomposition they describe the actual breaking-up of

the cells—the nuclei * fragmenting in the manner described by

Deineka for the Perch and the cytoplasm becoming converted

into the masses of granular matter found in the gland ducts and

bladder lumen. The principal reason which leads Nusbaum &

Reis to this surprising conclusion is the supposed incompetence

of the blood to supply the percentage of nitrogen gas found in the

bladder. Among minor features of this hypothesis of Nusbaum

& Reis, we may mention that they regard the disintegration of

the red blood corpuscles both as a source of nutrition for the gas

gland cells (the granular striping of the borders of gland cells in

contact with capillaries and the intracellular channels penetrating

the gland-cells being evidence of the cells absorbing this nutrition)

and as a source of the oxygen which, with the nutrition,

is urgently required by the cells in connection with their rapid

decomposition.

Thus the respective views of Jaeger and Nusbaum & Reis

largely differ as regards the function attributed to the individual

gas gland cell. Jaeger regards it as essentially a pumping

mechanism; N usbaum & Reis regard it as, primarily, mere

substance to be decomposed into its gaseous and solid constituents,

both being extruded into the bladder.

I will now proceed to discuss these two views as a preliminary

to a statement of my own views on the subject. Apart from

omissions common to both theories, which J shall endeavour to

remedy shortly, the chief fault to be found with the theory of

Jaeger is his refusal to recognize the origin of the gas bubbles in

the cells of the gas gland. Concerning the existence of these

* | may point out in this connection that during hemolysis the nucleus of

the red corpuscles appears to persist quite unchanged, even though the cytoplasm

may become entirely disintegrated.

GAS GLANDS OF SOME TELEOSTHAN FISHES. 225

structures there.can be no doubt whatever (see Appendix B),

They have been exhaustively studied by Bykowski and Nusbauin

& Reis, and [ have myself described and figured them in the

present paper. Jaeger himself, as [I have already stated, admits

the presence of vacuoles in the gas gland cells, but, curiously

enough, denies that they contain gas, and pronounces them

similar in nature to the vacuoles in the cells of the liver. It is

difficult to understand why this comparison should be made, since,

from his own standpoint, it is the function of the gas gland cells

to pump out gas and not to store up glycogen or fat. But even

adopting his suggestion that the supposed gas bubbles are only

vacuoles, we may remark that several authorities have described

in the kidney ‘the formation of vesicles in the cells and appear-

ances which indicate the discharge of these vesicles into the

cavity of the tubules” (37); and since kidney cells, like the gas

gland cells, are supposed by many modern physiologists to derive

most of their excreted substance by direct abstraction from the

blood, comparatively few of the constituents of urine being

Text-fig. 60 (x 1000).

—

Vacuoles in the kidney cells of Nerophis, some apparently being expelled into

the lumen of the tubule.

manufactured in the cell, we may certainly conclude that even

liquid vacuoles can be originated aud discharged in a manner

precisely comparable with that of the gas bubbles, and that if they

contained gas instead of liquid they would be indistinguishable.

I have figured some of the vacuoles present in the kidney cells of

the fish Nerophis (text-fig. 60). It is evident that in histological

preparations no sign of the liquid urine would be visible after

expulsion from the cell, whereas the gas bubble with its walls of

cytoplasmic material may, like a soap bubble in air, persist for

some time before bursting. Consistently with his denial of the

gaseous nature of the vacuoles in the gas cells, Jaeger was com-

pelled to regard the granular masses lying external to the gas

gland in the bladder lumen merely as a bye-product of the specific

activity of the gas gland.

If, however, we fully agree with Nusbaum & Reis as to the

226 DR. W. N. F. WOODLAND ON THE

origin of gas as bubbles in the cytoplasm of the gas gland cells,

we as certainly disagree with the interpretation they put upon

this phenomenon, viz. that the gas is produced as the result of

the actual chemical decomposition of the cytoplasm. Like Jaeger

we have found no evidence of that peculiar cell-disintegration

which is supposed by Nusbaum & Reis to be the histological

expression of the chemical decomposition of the cell. It is true,

as Jaeger admits, that in the cells of active gas glands the

cytoplasm (not the nucleus, which remains normal) often assumes

a “ hard-worked” appearance—the cytoplasm looks “ stringy” and

contains numerous empty spaces—and considering that the

process of continuous pumping of gas into the bladder must be

exceedingly arduous, this is not surprising. It is also true,

though not admitted by Jaeger, that the expulsion of bubbles

from the cell into the bladder involves, to a considerable extent,

waste of cell-substance. Each bubble, as already mentioned,

possesses a wall of cytoplasm, and on the bursting of the bubble

in the bladder lumen or gland duct, this wall, of course, breaks

down and contributes to the mass of granular matter found in the

bladder lumen and gland ducts. It must be confessed, therefore,

that cell-disintegration occurs to a considerable extent and is

associated with the production of bubbles of gas, but this

mechanical disintegration of cell-substance 1s quite another thing

from the hypothetical chemical decomposition of cell-substance

postulated by Nusbaum & Reis (see Appendix B). Jaeger

criticises this hypothesis of Nusbaum & Reis in a very effectual

manner. He first of all points out that it is impossible to regard

the epithelium of the gas gland as analogous in its mode of

working to a sebaceous gland, since the cells of the sebaceous

gland decompose in order to produce a highly-complex substance

chemically different from the substances supplied to them by the

blood, whereas the cells of the gas gland give rise to the most

simple of substances, viz. the gaseous elements oxygen and

nitrogen which are supplied to them ready-made. In other

words, the production of oxygen and nitrogen, unlike the secretion

of the sebaceous gland, requires no elaborate cell-metabolism, and

there is therefore no reason for the gas gland cell-decomposition

which Nusbaum & Reis affirm*. Further, according to Nusbaum

& Reis, the blood corpuscles in breaking up supply the gas gland

cells with oxygen for the decomposition of their suhstance, from

* We may, indeed, compare the process of gas-production with another process

familiar to the present writer, in which a simple substance is also supplied to the

cell and again liberated by that cell though in a different form, viz the deposition of

calcareous spicules in various invertebrate groups. Calcareous spicules have, like

gas bubbles, been regarded hy some investigators as resulting from the actual trans-

formation of cell-substance, but this view is not held by any modern zoologist. The

spicule-secreting cell is now regarded as a mechanisin for abstracting the dissolved

calcareous matter from the sea-water and of redepositing it, mixed with a variable

minute amount of organic substance, in a crystalline form, the aggregate of calcite

crystals constituting the spicule. The work of the cell in this case is simply the

abstraction and recrystallization of the dissolved calcareous salts—there 1s no question

of cell-decomposition. The mode of working of the kidney cell may also be compared

in this connection.

GAS GLANDS OF SOME TELEOSTEAN FISHES, DO

which decomposition oxygen is to be produced (!), but oxygen

cannot be used up in decomposing the gland cells and yet be

available to fill the bladder space. And as regards the supposed

production of free nitrogen by the decomposition of the gas gland

cells, Jaeger might have pointed out that such a fact would be

unique in animal metabolism. Free nitrogen is not known to be

liberated by any katabolic process. The only source of the free

nitrogen found in the bladder must be the small amount of that

gas dissolved in the blood plasma. But Jaeger’s chief objection

to the theory of Nusbaum & Reis is the impossibility of accounting

for the high pressure of the gas contained in the bladder, if this

gas be simply produced by the decomposition of the cell-substance.

The generation of carbon dioxide from chalk at once ceases, he

points out, despite a high temperature, if the gas be allowed to

accumulate in the calcination chamber, and in the same way the

decomposition of cytoplasm would cease long before the liberated

as could exert a fraction of the pressure usually found in the

bladder *. Oppel (57), in a summary of papers by Reis & Nusbaum,

also objects that whilst these authors strongly insist upon cell-

degeneration yet they have not stated in what way the epithelium

is regenerated after periods of activity, and I also have been

unable to come to a conclusion as to the opinion held by Reis &

Nusbaum on this subject. In one paper (62) these authors state

in the last two sentences that they have observed mitotic figures

in the gland cells, and conclude that regeneration of the gland is

effected by ordinary cell-division; in a subsequent paper (68),

however, they appear to embrace the curious view that the

amitosis first described by Deineka in Perca is a result of the

violent cell-decomposition associated, in their opinion, with the

activity of the gas gland, ¢. e. the nuclei, as well as the cytoplasm,

undergo disruptive changes, and that this amitosis (55) leads to

the production of new cells which take the place of those broken

up! Needless to say, I fail to see any justification for this

heenix-like theory. My own researches have proved that the

cells of the gas gland, when worn out like other cells of the body

by a long period of activity, are replaced by ordinary mitotic

divisions of the small cells at the base of the gland and that

amitosis is only found in the later cell-generations. I may also

mention in connection with this necrobiotic or decomposition

theory of Nusbaum & Reis that, apart from other objections, it

has always seemed to me to expect too much from the individual

cell of the gas gland. The cell is not only required to commit

suicide by undergoing decomposition but whilst undergoing

decomposition it is to work energetically + in pumping the results

of its own decay into the bladder cavity. Even the cell of the

228 DR. W. N. F. WOODLAND ON THE

sebaceous gland is not called upon for this degree of self-sacrifice.

Finally, the researches of Deineka (29) on the intimate nerve-

supply of the cells of the gas gland—it being stated that each

gland cell is surrounded by a terminal cluster of nerve-filaments—

and the known nervous control of the gas gland by two kinds of

fibres running in the vagus and sympathetic nerves respectively

(Moreau, 50, 51,52; Bohr, 20; and others) render it very im-

probable that such cells undergo the wasteful decomposition

assumed by Nusbaum & Reis.

I have now discussed the main features of the rival theories

of Jaeger and Nusbaum & Reis. If we eliminate the errors

and combine the truths of both with some observations of

other authorities, we arrive at the following outline statement

of the mode of working of the gas gland. The gas gland, in some

unexplained manner, but probably, as Jaeger suggests, by the

secretion of a toxin (which Reis, & Nusbaum suggest he should

have figured!) so influences the blood conveyed to it as to cause

the disintegration of a certain proportion of red blood corpuscles.

This fact of the disintegration is admitted by Bykowski and

Nusbaum & Reis. The disintegration of the erythrocytes

is primarily for the purpose of enabling the gas gland

cells to “lay hold” of the relatively large quantity of oxygen

contained in the blood. Oxygen, in consequence of its easy

obtainment from the blood, is the all-important gas required by

fishes which undergo considerable and rapid changes of pressure

in their bladder consequent on rapid changes in their vertical

position, and the gas gland, considered as a special development

of the lining epithelium of the bladder, is sclely concerned with

the rapid production of oxygen. The gas gland being essentially

an oxygen gland is therefore most developed in those fishes which

ossess the greatest proportion of oxygen in their bladder gas; in

tishes like the Cyprinide and some other freshwater families, on

the other hand—fishes which exist in inland waters possessing

little depth and which cannot, therefore, experience great changes

of pressure in their bladder by vertical displacement,—nitrogen

and carbon dioxide form the principal constituents of the bladder

gas * and gas glands, in the ordinary sense of the word, and retia

mirabilia are absent. It was formerly supposed that a large

proportion of oxygen in the bladder was associated with the great

depth at which the fish existed—the greater the depth the greater

the percentage of oxygen,—but apparently this is not altogether

the case f. A large proportion of oxygen, as already stated, seems

to be, for the most part, associated with the habits of those fish

which frequently make considerable excursions in a vertical

direction : if the fish sinks, the oxygen gland becomes active and

* Cyprinus carpio contains about 94 per cent. of nitrogen in its bladder (Htfner).

+ Ct. Hxocetus volitans, the Flying-fish, an essentially surface marine form,

possessing, according to Humboldt (41), 94 per cent. of nitrogen in its bladder, with

Coregonus acronius, a fresuwater form living permanently at a depth of over

seventy metres and possessing, according to Hiifner, about the same percentage of

nitrogen, and many other examples might be quoted.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 229

rapidly produces oxygen in considerable quantity in order to

counteract the compression of the bladder due to the increased

pressure and so to maintain the equality of the specific gravity of

the fish with its medium ; if the fish rises, either the absorption-

organ known as the “oval” comes into action and returns the

surplus oxygen in the swollen bladder to the blood, or, if the fish

be a physostome, the pneumatic duct allows the surplus gas to

escape to the exterior. Between the two extreme classes of

bladder—the oxygen-filled bladder with “red bodies” and the

nitrogen-filled bladder devoid of “red bodies”—there exist many

transitional kinds containing a relatively small percentage of

oxygen and feebly-developed ‘red bodies”*. As regards the

exact mode of abstraction by the gas gland of the gases contained

in the blood and their subsequent expulsion into the bladder,

papers already published afford little or no information on the

subject. In the case of those gases present in the bladder in

minute quantities, it is probable that they leave the blood by

simple diffusion: e.g. Traube-Mengarini (73, 74) and Priefer (60)

showed that this was the case when hydrogen was dissolved in the

water. It is evident, therefore, that the problem of gas-

abstraction becomes of importance only when the partial pressure

of any particular gas in the bladder exceeds that in the blood.

a—TT . . 1 . .

That the gas gland cells act as.a pump is certain t; it is also

certain that the gases first enter the cells in a dissolved condition

and that, just as a scleroblast converts dissolved calcareous salts

into a solid spicule,/so the gas gland cells cause these dissolved

gases to appear in 4 gaseous form as bubbles in the cytoplasm §.

In addition to this, however, the gland cells subject these gas

bubbles to a considerable pressure, greater than that existing in

the bladder, so that when the gas bubbles are expelled into the

bladder lumen and experience a diminution of pressure they

burst. As regards the varying composition of the bladder gas, it

can only be concluded that the cells of the gas gland, like the cells

of the kidney, exercise a selective power. That the oxygen

pumped into the bladder is derived from the blood no one doubts,

but Nusbaum & Reis, Thilo, and some others find a great

difficulty im supposing that nitrogen (and carbon dioxide) is

* Hig. Perca fluviatilis contains on an average 15 per cent. of oxygen, 83 of nitrogen,

and 2 of carbon dioxide (Hiifner) ; Lota vulgaris 65 per cent. oxygen, 30 nitrogen,

5 carbon dioxide (Hiitner), &c., &c. For analyses of the bladder gases in many

fish see the works of Biot (19), Configliachi (27), Delaroche (80), Humboldt &

Provengal (89, 40), Hiifner (88), Richard (6'7) among others. The percentage

composition of the bladder gas exhibits, as might be expected, considerable variation

not only in different individuals of the same species of fish but in the same individual

at different times.

+ It must be mentioned, however, that Humboldt & Provencal (89) in 1809 per-

formed this experiment of impregnating water with hydrogen and failed to detect a

trace of hydrogen in the bladder, but probably their methods of analysis were too

ciude.

+ In this connection Hiifner ventures to compare the gas gland cells with the

contractile gland cells described by Drasch (82) in the neck-skin of the Frog, both

being similarly supplied with nerves.

§ Cf. the gas vacuoles in Arcella and other Thecamebida.

Proc. Zoou. Soc.—1911, No. XVI. 16

230 DR. W. N. F. WOODLAND ON THE

similarly derived. It is true that the amount of nitrogen

dissolved in the blood is very minute as compared with the

amount of oxygen, but, on the other hand, nitrogen does not, like

oxygen, require to be produced at a rapid rate; and since the

supposition of Nusbaum & Reis, that free nitrogen is generated

by the decomposition of cell-substance, is quite inadmissible for

the reasons supplied above, the nitrogen dissolved in the blood

can be the only source of that gas. It is well known that human

blood, when suddenly released from great pressure, develops

bubbles of nitrogen owing to the inability of the blood to re-

dissolve the gas immediately, and doubtless the cells of the gas

gland also so act upon (though certainly not by a diminution of

pressure) the blood plasma bathing their substance as to compel

the nitrogen to assume a gaseous form. Once abstracted from

the blood stream, the nitrogen, in the case of those bladders

containing nitrogen at high pressure, is pumped in the same

way as, and in many cases with, the oxygen into the bladder

lumen *.

I now propose to offer some additional suggestions concerning

the physiology of the gas gland which have occurred to me

during the course of my work. It is curious that not one of the

investigators mentioned in the foregoing pages has attempted to

explain in a satisfactory manner the striking conformation of the

rete mirabile. A year ago I published in a short note (78)

(based, as I have already explained, upon a mistaken conception

of the teleost pancre as) a sketch of a new theory concerning

the use of the rete which I have again outlined on a preceding

page of this paper, but apart from ‘this I have met with no

suggestion concerning the physiological significance of the rete

mirabile bipolare geminum (or more simply rete mirabile duplex)

I now propose to restate more fully this theory of mine, but

before doing so I will mention the only previous hypothesis of

which I am aware. Johannes Miiller (53) stated that in his

opinion the retia mirabilia associated with the gas glands of

teleost fishes (and Miller was one of the first, if not the first, to

distinguish the gas glands—“ luftdriisen”—from the retia mira-

bilia and to state their proper function as gas-producers) possessed

the same utility as the various other kinds of retia mirabilia de-

scribed by him, viz. to cause the blood stream to flow more slowly

for some physiological purpose, and this opinion has been adopted,

so far as I know, by all subsequent observers. Presumably

the slowness of the blood stream in connection with the gas glands

is supposed to be for the purpose of allowing the cells of the

gas gland time in which to abstract the gases present in the blood,

and that this is one function of the rete: mirabile I myself do

* Haldane (86) comes to the same conclusion.

+ The following suggestion concerning the use of the duplex rete associated with

the gas glands may throw some light upon the physiology of the similar duplex

retia mirabilia which are stated to occur in connection with other structures,

e.g. the “choroid gland” associated with the eye of many teleosts and the rete

im connection with the liver of the Tunny.

GAS GLANDS OF SOME TELBOSTEAN FISHES. 231

not doubt.* But thisexplanation does not, in my opinion, explain

all the facts, since, if mere slowing down of the blood current

is the one desideratum, then a rete solely connected with the

bladder artery is all-sufficient. Further, it is evident that the

explanation takes no account either of the fact that the bladder

vein also forms a rete mirabile or of the still more remarkable fact

that the retia on the bladder artery and vein respectively are both

Jormed at exactly the same distance from the gas gland and their

constituent arterioles and venules as intimately intermingled as any

product of human manufacture purposely so designed could be, The

suggestion which I venture to offer as to the use of this complex

arterial and venous rete universally associated with gas glands

is the natural conclusion of the following considerations, The

fact that some hundreds of the finest capillaries conveying

blood ¢o the gas gland are intimately intermingled with and

closely apposed to a like number (another significant fact) of

similar capillaries conveying blood from the gas gland is sug-

gestive of the exchange between the two sets of capillaries of some

substance necessarily of importance to the gas gland with which

the rete is connected. This hypothetical substance we must

assume both to be diffused from the venous capillaries to the

arterial (since diffusion in an opposite direction would not affect

the gas gland) and to be derived from the gas gland. The

question which next arises is why this hypothetical substance

should be poured into the arterial blood before it reaches the

gas gland, and the only answer that suggests itself is that

it is necessary for this substance to influence the arterial blood

in some manner during its passage (made slower by the rete)

from the rete to the gas gland, so that by the time the arterial

blood reaches its destination, its constitution has become altered.

Now I have already stated that the gas gland, in the limited

sense of the term, is essentially an oxygen-producing gland and

that therefore the oxygen contained in the blood is the one

element which the gas gland requires, from which fact we may

conclude that the hypothetical substance referred to has some-

thing to do with the giving up of oxygen by the blood to the

gland. In the foregoing lines I have mentioned that such a

hypothetical substance has already been postulated by Jaeger to

explain the disintegration of the red blood corpuscles. We may

therefore state as extremely probable suppositions that in the rete

the venous capillaries contain a relatively large quantity of a

toxin poured into the blood by the cells of the gas gland, that

this toxin diffuses from the venous capillaries into the arterial,

and that whilst the arterial blood is slowly travelling from the

* This function, however, cannot he exercised in the case of the retia mirabilia of

the Eel, which are separated from the gas gland by a relatively few large vessels,

since the reunion of the arterial capillaries on the side next the gas gland means of

course the quickening of the blood stream. Neither can the Hel’s rete mirabile

possess the supposed function of the “ carotid gland” of Amphibia, of “ deadening ”

the heart pulse, since the gill capillary system of the fish must effectively eliminate

all trace of this; on the other hand, the rete mirabile must cervainly minimize the

pressure of the blood supplying the gas gland, a result which, in the present instance,

we must regard as a defect,

16*

232 DR. W. N. F. WOODLAND ON THE

rete to the gland the toxin eftects by a process of haemolysis the

partial or total dismtegration of a certain proportion of erythro-

cytes, the cytoplasm and oxyhzmoglobin pigment of each being

scattered in the plasma, the former giving rise to the masses of

granular matter so largely present in the blood returned from the

bladder and the latter becoming dissolved in the plasma.

Is there any evidence of this process actually taking place ?

In reply to this question I am able to cite one fact, already stated

on page 219 in Part I., which I think constitutes strong evidence in

the affirmative. It is evident that if this process takes place

then we may expect to find at the proximal pole of the rete, 7. e.

the pole remote from the gas gland, large quantities of granular

matter (resulting from the erythrocytolysis above described)

present in the venous capillaries but little or none in the arterial,

and that towards the distal pole of the rete (the toxin, on the

hypothesis, having had time to diffuse from the venous capillaries

and influence the arterial blood) granular débris will also appear

in the arteries. Careful examination of the rete mirabile asso-

ciated with active glands has convinced me that this inferred

distribution of the granular matter in the rete capillaries is a fact.

Figure 32 (Pl. IV.) represents a section taken acrcss the proximal

pole of the rete and shows the absence of the erythrocytolytic

granules in the arteries; a section across the distal end of the rete,

on the other hand, shows that hemolysis of the arterial blood has

commenced, erythrocytolytic granules being present.

Oppel (56) puts a question which will be asked by many other

critics of the foregoing theory of “red body” physiology. Why,

he asks, is it at all necessary for the erythrocytes to be broken up

in order that oxygen may be supplied to the cells of the gas gland @

In other words, if, as we kuow is the case, the blood readily gives

up its oxygen to supply the ordinary tissues of the body, where

is the necessity for the manufacture of a special toxic substance

in order to effect this same liberation of oxygen in the case of the

gas gland? Jam not aware that Jaeger or any other author has

given a definite answer to this question, but a consideration of

the facts will soon supply one. In the first place, it is evident

that the cells of the gas gland stand in a very different relation to

the oxygen of the blood as compared with that of ordinary tissue

cells, since whereas the latter are deficient in oxygen and require

it solely for purposes of metabolism, the former are already

saturated with oxygen and only lay hold of it in order to con-

centrate it and pump it into the bladder; in other words, the

cells of the gas gland require to get a very effectual “ grip” on

the oxygen which the tissue cells do not *. Secondly, realization

of the conditions obtaining in the normal supply of oxygen by

the blood to the tissues will enable us to understand the necessity

for a toxin. The combination of oxygen with the hemoglobin

* This necessity for the gas gland cells to be in the closest contact with the bicod

is possibly accountable for the presence of the intracellular capillaries already

described, perhaps also for the intracellular lumina as conveyors of exuded plasma

into the celi-substance. The same phenomena are met with in the cells of the

liver—an organ which also requires close contact with the blood—, the canaliculi

representing the intracellular channels.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 233

pigment contained in the red corpuscles is, as is well known, a

very loose one, and this loose combination is, according to modern

views, only maintained by the maximum partial pressure exerted

by the small amount of oxygen dissolved in the blood plasma.

Further, it is important to notice in connection with the present

subject that in the liberation of oxygen to the tissues of the body,

it is the oxygen dissolved in the plasma which is immediately

supplied to the tissues, and not the oxygen associated with the

hemoglobin. The oxygen of the oxyhemoglobin compound only

becomes liberated as a consequence of the lowering of the partial

pressure of the plasma oxygen caused by tissue absorption and

when liberated merely serves to replenish the plasma and is not

conveyed directly to the tissues, so that the plasma with its

limited solvent action on oxygen must be recognized as the all-

important intermediary between the store of oxygen combined

with the hemoglobin and the tissues. Now in the case of the

cells of the gas gland, already employed in pumping oxygen into

the bladder and certainly possessing all and more than they

require for metabolic purposes, it is difficult to imagine that they

can absorb the oxygen dissolved in the plasma in the manner

employed by ordinary tissue cells. Even if they are so able, the

small amount of oxygen so obtainable is quite insufficient for

their purposes, since the oxygen, unlike the nitrogen and carbon

dioxide, is required to be rapidly produced in large quantities.

The only alternative is, then, for the gas gland cells to seize oa

the main source of the oxygen, viz. the oxyhemoglobin, and s

obtain in wholesale quantity what the plasma can only supply in

retail*. This is effected, as we have seen, by the production of a

toxin which, doubtless by a process of hemolysis, breaks up the

red corpuscles into fragments and so liberates the contained

oxyhzemoglobin into the plasma, the rete ensuring that this

hemolysis and consequent solution of oxyhemoglobin in the

plasma i is effected in time for the dissolved pigment to be available

* This reduction of the blood to the primitive invertebrate condition in which the

respivatory pigment is dissolved in the general plasma and not imprisoned in elastic

discs (the erythrocytes) as in Vertebrates raises the question as to why, if the former

condition enables the tissues to absorb the oxygen more readily, the latter condition

has arisen. Apart from a few lamellibranch ‘and other Mollusca and a few Poly-

chietes and Phoronis in which hematids have been described, all Invertebrates (and,

according to Lankester, also Amphioxus and the Leptocephalus larva of the Hel, but

hzematids have been stated to occur in the former and possibly exist in small number

in the latter) carry the respiratory pigment, when this is present (absent in the

tracheate. Arthropods, e. g.), in the plasma. and it is difficult to understand what

advantage accrues from preventing the dissolved oxyhemoglobin coming into direct

contact with the tissues in vertebrate animals. It is of course possible that the

indirect distribution of oxygen by way of solution in the plasma conduces to a more

even and gradual supply to the tissues, especially in animals like Vertebrates in

which most of the tissues are very remote from the limited respiratory area of the

body surface; in most Invertebrates, on the other hand, oxidation of the blood takes

place over most of the body surface and the tissues are all practically simultaneously

reached by the blood which, being contained in sinusoids rather than in capillaries,

bathes them on all sides. The more vapid circulation of the blood and larger

quantity of hemoglobin in Vertebrates possibly compensate for the absence of

hemoglobin in the plasma. As is well known, when haemoglobin is liberated into

the plasma in Vertebrate blood, it is at once eliminated by the liver and kidneys.

See Addenda (2).

234 DR. W. N. F. WOODLAND ON THE

for absorption when the blood reaches the gas gland. Is there

any evidence that the ceils of the gas gland absorb this oxy-

hemoglobin dissolved in the blood plasma? It is not too much

to say in reply that every preparation of a gas gland in at all

an active condition does provide very substantial evidence as to

the actual occurrence of this process. In every such preparation

(see figs. 28-31, 50, 68, 73, e. g.) it is at once noticeable that the

cytoplasm of the gas gland cells in contact with the capillaries-—

and usually the cells are only separated from the blood by the

thin endothelium—is of dense appearance and forms a perivascular

zone quite distinct from the rest of the ceil-protoplasm, a feature

which has been described and figured by all recent observers of

gas gland structure—Jaeger, Bykowski, Nusbaum, Reis and the

present writer (see Part I.). Nusbaum & Reis hold that this

appearance of the cytoplasm next the blood vessels is merely evi-

dence of absorption fromthe blood of the nutritious matter afforded

by the disintegration of the erythrocytes, but for the reasons just

given we prefer to believe that the oxygen associated with the

hemoglobin is éhe desideratum of the gland cells and that what-

ever nutritive value ingested fragments of stroma may possess 1S

quite a minor matter. However, the opinion of Nusbaum &

Reis is of value in supporting our conclusion that the cells of the

gas gland do actually absorb from the blood material liberated by

the breaking-up of the erythrocytes. Hxamination of good

preparations of active glands shows that this darkening of the

cytoplasm of the individual gas gland cell situated next the blood-

channel really possesses a striped appearance (Pl. IX. fig. 73)—

Bykowski & Nusbaum, e. g., describe it as ‘die charakterische

Streifung des Protoplasmas rings um die Blutgefasse ”—similar to

that seen at the edges of the cells lining portions of the gut, in

the Sertoli cells of the testis and in other cases, and this striping

found in so many kinds of cells is proof of a process of absorption

taking place. We do not suppose that the fragments of corpuscle

substance composing the granular matter in the blood stream are

absorbed by the gas gland cells, but only the oxyhemoglobin

dissolved in the plasma, and the fact, which we have previously

stated, that the veins of the rete mirabile are in active glands full

of this granular matter is in accordance with this view. It can

thus be proved by what practically amounts to actual demon-

stration that the cells of the gas gland do absorb the dissolved

oxyhemoglobin directly from the blood (see Addenda (3)), and the

natural inference is that this is employed for the supply of oxygen

to the bladder.

Judging from analogies provided by other classes of secreting

cells, what possibly happens in the metabolism of the gas gland

cell in the production of bubbles of oxygen gas from the absorbed

oxyhemoglobin dissolved in the blood plasma is that at that end or

pole of the gas gland cell situated next the blood stream the

dissolved oxyhemoglobin forms a loose combination with the

cytoplasm, this combination being merely a temporary linkage of

the molecules of the two substances (similar, e. g., to the linkage of

GAS GLANDS OF SOME TELEOSTEAN FISHES. | 935

side-chains forming ‘“ anti-bodies,” which Ehrlich assumes to

occur in his theory of immunity) and that later at the remote

pole of the cell, i. e. the region of the cell remote from the blood

stream, this loose temporary incorporation of the dissolved

oxyhemoglobin with the cytoplasm breaks down with the

liberation of oxygen gas, which, as already seen, arises as bubbles

usually in the vicinity of the nucleus. The slight decomposition

of cell-substance which we may thus suppose to occur is evidently

quite distinct from that decomposition postulated by Nusbaum

& Reis. In our supposition the temporary linkage of the

oxyhemoglobin with the cytoplasm is merely to enable the

cytoplasm to obtain a “ grip” on the oxyhemoglobin in order to

dissociate the oxygen; in the supposition of ‘Nusbaum & Reis

it is the cytoplasm itself which decomposes.

A minor point remains to be mentioned. We have already

stated that the veins returning the blood from the gas gland

contain a relatively large amount of granular matter resulting

from the hemolysis of the red blood corpuscles, and the question

remains as to what becomes of this superfluous granular matter.

It can only be said in reply that probably a large portion of this

disintegration material is eliminated from the blood by the liver,

since we know that one function of the liver cells in all Verte-

brates is to destroy degenerate corpuscles* and other waste

material in the blood; possibly also the spleen assists in this

connection, though, judging from its histological appearance, I

doubt it. I have also observed in several genera (Verophis,

Clobius, Gasterosteus, &c.) masses of cells lying to the outer sides

of or between the kidneys in which a destruction of effete blood-

corpuscles seems to be actively proceeding. It is, however, of

little concern to us in what manner the blood is clarified in these

teleostean fishes ; it suffices to say that the relative purity of the

arterial blood in the biadder proves that such elimination does

take place.

On reviewing the foregoing pages it will be seen that the hypo-

thesis concerning the physiology of the “red bodies ” just elaborated

has much to be said for it. In its essential features it is the theory

of Jaeger and Hiifner, supplemented, however, and in some respects

corr ected, by the Coenan: of Bykowski, Nepean Reis, and

other investigators, and slightly extended by the few suggestions

made by the present writer. Its validity is assured by the number

of diverse facts which it interprets, for not only does it explain

the general fact of the existence of the “red body” as the only

mechanism possible under the conditions for the rapid inflation

of the bladder, but it also explains the minutest details of this

mechanism, such as the perivascular striping of the component

cells of the gas gland and the disintegration of the erythrocytes,

and such a wide range of interpretation constitutes the criterion

of a true theory.

* These erythrocytes in the liver have of course been largely deprived of their

oxygen by the gut tissues.

236 DR. W. N. F. WOODLAND ON THE

APPENDIX A.

The Sources and Modes of Preparation of Material.

A great part of my material was obtained by me during my occu-

pation of the British Association Table at the Naples Zoological

Station during April, 1907; additional specimens were sub-

sequently sent to me from Naples and the remainder I obtained

from the Plymouth Marine Biological Station. As regards

methods of preparation, I may remark, first of all, that it is

important to fix the lining-epithelium of the bladder and its

special development, the gas gland, in a distended condition, and

this is best effected by puncturing the bladder (in sitw) posteriorly

and immediately filling it with the fixative used, also pouring

fixative on the outside of the bladder. By this means all shrinkage

of the bladder wall is avoided on removal fromthe body. Another

advisable precaution is to separate the lining-epithelium of the

bladder, after it has been well fixed, from the outer layers of the

bladder wall. since I found that in many cases the tough muscle

and connective tissue composing these latter cause difficulty in

obtaining thin sections across the gas gland. The fixatives which

IT employed chiefly were Zenker’s Fluid, Corrosive Acetic, and

Mann’s Fluid (Distilled Water 100 c.c., Corros. Sub. 2°5 gm., Picric

Acid 1 gm.. 40°/, Formol 10-25 ¢.c.—a fixative which, in conjunc-

tion with Borax Carmine and Picro-indigo-carmine, gave me most

beautiful results). Some of my specimens were simply fixed in

10 °/, Formol—a fixative which did not yield such good results as

the others, though good enough for most purposes. I left objects

in Zenker and Mann overnight, but in Corrosive Acetic only for

an hour or so. In many cases I fixed duplicate specimens in dif-

ferent fixatives and was thus able to compare results.g Objects

fixed with Zenker were well washed for several hours with Distilled

Water and then, like the remaining objects fixed with other fix-

atives, graded slowly up to 70°/, Alcohol (the percentages being

in all cases made up with Absolute Alcohol and Distilled Water).

Objects fixed with fluids containing Corrosive Sublimate were

treated with Iodine at this stage. In most cases I subsequently

stained in bulk with Grenacher’s Borax Carmine, leaving the

objects in for at least twelve hours, and without differentiation with

Acid Alcohol, dehydrated, cleared,and embedded in Paraffin. The

sections (both longitudinal and transverse in many cases), well

stained with Borax Carmine, were then stained on the slide with

Picro-indigo-carmine *. This stain is made up by adding one part

of a saturated solution of Picric Acid in 90°/, Aleohel (sat. sol.

=cirea 4:5°/,) to two parts of a saturated solution of Grubler’s

Indig-carmin in 70°/, Alcohol (sat. sol.=circa 1°/,), and it is well

to dilute this stain so obtained with twice its bulk of 70°/, Alcohol.

The sections are placed in this stain for a short time, varying

according to the thickness of the sections and depth of staining

with Borax Carmine, from a few minutes to over half an hour.

* T am indebted to Mr. A. D. Darbishire for recommending me this stain.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 237

The Picric Acid, of course, differentiates the Borax Carmine, leaving

scarlet nuclei, and the Picro-indigo-carmine stains the cytoplasm

of the gas gland a dull green, the cytoplasm of red blood corpuscles

and secretion products like zymogen granules a brilliant emerald-

green, and connective tissue blue. Curiously enough, I did not

always obtain these brilliant colour-contrasts with Zenker, espe-

cially if the material had been preserved in Alcohol and Glycerine

for some time after fixation, but fixation with Mann’s Fluid always

yielded the best results. Objects which did not stain well with

the Borax Carmine [ stained with Ehrlich’s Hematoxylin and

Picro-indigo-carmine, and obtained sufficiently good contrasts. 'The

principal object to be attained by using Picro-indigo-carmine is to

show up the blood corpuscles and granules. In exceptional cases

I employed the Ivon-hematoxylin method for nuclear details (in

Perca, e.g.), also Kernschwarz for rendering visible cytoplasmic

edges and hence the limits of intercellular and intracellular ducts

(in Gadus, e.g.). As above mentioned, most of my gas glands were

eut both transversely and longitudinally.

APPENDIX B.

On the Artificial Production of Gas Bubbles in Cells of the

Gas Gland.

Of the various methods available for compelling fish to produce

gas in their bladder I have employed several, but only with one

have I succeeded in detecting the gas bubbles in the act of being

formed by the cells. For the guidance of others I will first men-

tion my unsuccessful experiments. I first experimented with

Gobwus paganellus at Plymouth, subjecting this fish to increased

pressure due to increased depth—one of the easiest methods of

inducing gas-production. I took tank specimens from the Biolo-

gical Laboratory on board the steam launch, and when anchored

well out at sea I enclosed three or four specimens in each of five

cages. One cage I let down to a depth of 30 feet and kept it

there for one hour and a half; a second cage I let down 60 feet

for one hour, a third 90 feet for one hour, a fourth 180 feet for

half an hour and a fifth 180 feet for two and a half hours. On

examining sections of the gas glands of these fish I could detect

no decisive differences in the gland cells. J am unable to explain

why this experiment was a failure. Possibly even the maximum

time allowed (22 hours) was too short in which to allow these cells

to become active; possibly also the fact that Gobius paganellus is

a bottom form may have contributed to this negative result.

Another experiment which I may mention was to stimulate the

two vagus nerves of an Kel (Anguwilla\ with a battery for three

hours, the two sympathetics having been cut and the bladder

previously emptied and ligatured. At the end of the three hours

the bladder certainly contained about one-sixth of its usual volume

of gas, but I was unable to detect bubbles in the cytoplasm. I did

_not try the experiment of injecting fish with Pilocarpine nitrate

or any similar drug.

238 DR. W. N. F. WOODLAND ON YHE

The method by which I achieved success was that of attaching

weights tosome Perch ina tank in the manner described by Moreau*.

The fish, if appropriately weighted, increase the volume of gas in

their bladders in order to counteract the sinking action of the

weights, but, according to my experience, it is all-important that

the weights should not be too heavy, since if the fish are perma-

nently attached to the bottom—if there is no hope of being able to

rise—they apparently make no attempt at gas-production. The

form of weight [employed was a half-hoop of lead weighing about

5 gm. (approximately one-tenth the weight of the fish, and this

was probably too heavy); this I suspended round the lower half of

the fish (by thin wires tied dorsally) in the manner of a belt

passing dorso-ventrally between the pectoral and pelvic fins, so

leaving these perfectly free, After some twelve hours or more

the fish can swim about more easily and with a lighter weight it

would regain all its usual mobility. If now the weight be removed

the fish willrise quite helplessly to the surface of the water, owing

to the increased quantity of gas in the bladder. Unless the fish

so rises after the experiment 1t is of little use to examine the gas

gland. It is also, of course, necessary to compare the gas gland

sections of the fish experimented on with similar sections of a

control fish in the same tank. I weighted some six or seven Perch

for 17 hours and at the end of that period the bladder of each was

appreciably swollen, the fish having to swim vigorously in order to

keep below the surface after removal of the weight. I emptied

the bladder in the usual manner by a small puncture posteriorly

and in several cases refilled it with Zenker’s Fluid; in others I

used a mixture of one volume of Glacial Acetic with two volumes

of Absolute Alcohol, and in others 1°/,Osmic Acid. Bladders

fixed with Zenker I stained with Ehrlich and Picro-indigo-car-

mine; bladders fixed with the Glacial-Absolute mixture I stained

with Jron-haematoxylin in order to study the details of nuclear

structure; bladders fixed with Osmic I stained with Ranvier’s

Picro-carmine. In all cases I made surface-view preparations,

laying the bladder with its gas gland inside uppermost on the

slide, in addition to cutting numerous sections of various thick-

nesses of the Zenker and Glacial-Absolute material. The Zenker

material gave the best results from the gas bubble standpoint,

probably owing to the absence of strong diffusion-currents during

the process of fixation ; the Glacial-A bsolute mixture, on the other

hand, though perfect in its fixation of most parts of the cell, was

doubtless somewhat drastic in its action on delicate structures lke

extruded gas bubbles.

The sections of the Zenker material showed up the production

of the gas bubbles in a very effective manner, the bubbles being

clearly seen to arise in the cell-substance (not shown in text-fig. 61,

p- 240) and to be extruded as foam-like masses all over the surface

of the gas gland lining the bladder (Pl. 1X. fig. 70 and text-fig. 61,

the latter being a microphotograph). It must be remarked that

only occasional cells were thus active, the majority not exhibiting

* A more simple method, of course, is to empty the bladder with a trocar.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 939

bubbles. Gas glands must be in a very active condition for the

majority of cells to exhibit bubble-stcuctures (see Pl. V. fig. 35 of

Gobius minutus). These bubbles were quite absent in the sections

of non-weighted Perch used for the purpose of controlling the

experiment. In my Glacial-Absolute material, structures which

are presumably bubbles (very unlike the bubbles seen in most

preparations, however) are also to be found in the act of extru-

sion, but these instead of being unmistakable foam-like masses

are bladder-like structures projecting from the cell-periphery

(Pl. IX. fig. 71 and text-fig. 62, the latter being a microphoto-

graph) and some are to be found lying free in the bladder-cavity.

Possibly the action of the fixative is to cause all the small bubbles

of the foam-like mass to run together to form one big bubble.

In the surface-view preparations fixed by this method, large gas

bubbles are occasionally to be found in the cytoplasm. All these

bubble-structures are quite absent in the control material fixed

with Glacial-Absolute.

Apart from the existence of gas bubbles in preparations of the

activated gland, another distinction from the non-active or dormant

gland is the “worked” or ‘‘spent” appearance of many of the cells,

a distinction which I have tried to indicate in figures 71 and 72

(71 represents cells of the active gland—c/. text-fig.62; 72 repre-

sents dormant cells). It was this “spent” appearance, this

attenuated and ragged condition of the cytoplasm, which suggested

to Nusbaum & Reis that the cell substance actually decomposed

into the bladder gases and granular masses, just as the cells of the

sebaceous gland disintegrate to produce the secretion of that gland,

and, indeed, at first sight of these exhausted cells of the gas gland

the suggestion does not seem unreasonable. However, the fact.

that a cell which posseses but a small amount of protoplasm in the

space bounded by its walls is not necessarily in a state of decom-

position is shown by the majority of plant-cells, the cytoplasm

of which is reduced to a “ primordial utricle,” by swollen adipose

cells, by spicule-cells and by yolk-laden meroblastic eggs, to men-

tion a few out of many possible examples, and since no amount of

cytological technique can demonstrate the actual chemical decom-

position Nusbaum & Reis assume, this attenuation of the

substance of the cells of the gas gland is not of very great

significance, especially in view of the reasons for rejecting this

suggestion of Nusbaum & Reis already advanced in Part IJ. It

must not be supposed that the cells of my activated glands alone

wear this ‘‘spent” aspect, all the cells of the dormant gland

resembling figure 72; on the contrary, the only difference between

the cells of the two glands is one of degree, cells of the dormant

gland being, on the whole, less ragged than those of the active.

Only a gas gland completely at rest (a condition probably rarely

attained in a voracious fish like the Perch) would have all its cells

in a resting condition. In most marine fish, however, which I

have studied, nearly all the cells of the resting gland have been

quite solid in appearance, only the cells of the active gland being

attenuated.

240, DR. W. N. F. WOODLAND ON THE

Text-fig. 61 (x cire. 1000).

Micrcphotograph of transverse section across thin region of gas gland of Perea (fixed

with Zenker’s Fluid), showing foam-like mass of bubbles (F.B.) on surface

(of. Pl. IX. fig. 70).

Text-fig. 62 (x cire. 1000).

Microphotograph of transverse section across thick region of gas gland of Perea (fixed

with Absolute and Glacial Acetic), showing extruded gas bubbles (B) on surface

(cf. Pl. IX. fig. 71). The “spent” condition of the cells is also shown.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 241

In connection with the two microphotographs of gas bubbles

here reproduced as text-figures 61 and 62, I may say that these

structures were in my preparations very difficult subjects for

photography, partly because of the inappropriate staining em-

ployed and partly because bubbles shown in optical section of

necessity bear but little resemblance to the real thing (this was

especially the case with the foam-like mass indicated in text-fig. 61 ;

cf. fig. 70). Nevertheless, these microphotographs, taken for me by

Mr. F.J. Pittock, of the Zoological Department, University College,

are perhaps of value as affording impartial evidence of the

appearance of an active gas gland epithelium, also of the ‘“ spent ”

condition of the cytoplasm. Microphotographs were also taken of

intracellular bubbles, but, owing to the thickness and staining of

the sections, were not suitable for reproduction.

ADDENDA.

(1) The statement that gas glands are normally quiescent may

not be strictly accurate ; it would perhaps be more correct to say

that only on occasion do gas glands assume great activity. I

make this remark in consequence of a communication from

Lieutenant G. C. C. Damant, R.N., who kindly permits me to

repeat it. From observations made under water during diving

operations and from the results obtained in catching fish on the

hook at different states of the tide, Lieut. Damant concludes that

Pout, e. g., remain fairly constantly at one level, viz., just off the

bottom, whatever the state of the tide may be. If this be the

case, it follows, as Lieut. Damant suggests, that the gas gland

must become functional during each rise of the tide in order that

a periodic increase of gas in the bladder may counteract the

periodic increase of external pressure.

(2) It seems probable, from suggestions kindly made to me by

Dr. G. A. Buckmaster and Dr. G. C. Mathison, that the principal

factor in the evolution of erythrocytes in Vertebrates has been

the necessity for a greatly increased quantity of hemoglobin in

Vertebrate blood as compared with Invertebrate—a necessity

corresponding to greater functional activity. Itis suggested that

the amount of hemoglobin required to be present in Vertebrate

blood would be more than the plasma could possibly hold in

solution, and that even if this were possible the viscosity of the

plasma produced would seriously interfere with the other func-

tions of the plasma, which, like the respiratory, have, in

Vertebrates, increased in intensity. This large quantity of hemo-

globin in Vertebrate blood being requisite and its presence in the

plasma prohibited, the evolution of erythrocytes in which the

hemoglobin is imprisoned and combined with a proteid has

apparently been the only alternative. As I have before remarked,

the lack of direct contact between the tissues and the respiratory

pigment in Vertebrates is compensated for both by the rapidity of

the circulation and by the amount of the respiratory pigment

present, and doubtless also by the increase in amount of the waste

242, DR. W. N. F. WOODLAND ON THE

products of active tissues which act as reducing agents and so

facilitate the abstraction of the oxygen dissolved in the plasma.

-(3) I have adduced the striped darkening of the cytoplasm of

the gas gland cells bordering the capillaries as evidence of the

absorption of dissolved oxyhemoglobin from the blood and that

this is evidence of absorption is undoubted, but that it is absorp-

tion of oxyhemoglobin is of course only an inference from the

hemolysis of the blood corpuscles, the production of oxygen gas

by the gland cells and other considerations. I wish to state

here, previous to sending away the final proofs of this paper, that,

judging from preliminary experiments with the Israel-Pappen-

heim stain for hemoglobin, kindly recommended to me by

Dr. G. A. Buckmaster, I have been able to detect the actual

presence of hemoglobin in the cells of the active gas gland and

its passage thereto from the blood stream, and Dr. Buckmaster

has confirmed these preliminary observations of mine. I intend

to repeat these observations on an extended scale on fresh

material shortly, and until I have done so I do not wish the

above statement to be considered as final.

References to Pancreas Literature (pp. 198, 200 in text).

1. Date, H. H.—<On the Islets of Langerhans in the Pancreas.” Phil. Trans.

Roy. Soc. Lond., Vol. excvil, 1905, pp. 25-46.

2, pp Wirt, L. M.—* The Morphology and Physiology of Areas of Langerhans in

some Vertebrates.” Jour. Exper. Med. New York, Vol. vin, 1906,

pp. 193-239.

3. Dramare, V-—“Studii comparativi sulle isole di Langerhans de paneréas.”

Intern. Monatsschr. f. Anat. u. Physiol. Leipzig, Bd. xvi, 1899,

pp. 155-209.

3a. Furnt, M—* Das Bindgewebe der Speicheldrtisen und der Pankreas, ete.”

Aych. f. Anat. u. Entwick. Leipzig, Jahrg. 1903, pp. 61-106.

4. Lacuesse, H.-—“ Développement du Paneréas chez les Poissons Osseux.”

Comptes rendus hebdom. des Séances et Mémoires de la Soc. de

Biologie, Sér. 9, T. 1, 1889, p. 341.

———. “Pancréas intra-hépatique chez les Poissons.” Op. cié., Sér. 9, T. ui,

1891, p. 145.

——— “Surla Formation des ilots de Langerhans dans le Pancréas.’

Sér. 9, T. v, 1893, pp. 819, 820.

“ Développement du Pancréas chez Jes Poissons Osseux.” Journal

de Anatomie et Physiologie, Paris, T. xxx, 1894, p. 79.

———._ “Structure et Développement du Pancréas d/aprés les travaux récents.””

Thid. pp. 591-608.

———._ “Sur le Pancréas du Crénilabre et particuliérement sur le Pancréas

intrahépatique.”’ Revue Biologique du Nord de la France, T. vii,

1894-1895, pp. 343-360.

10. ——— “Sur la Structure du Pancréas chez quelques Ophidiens et particuli-

érement sur les tlots endocrimes.” Arch. d’Anat. Micr. Paris, T. iv,

1901, pp. 157-218.

11. Lays, M. A.—* The Cytological Characters of the Areas of Langerhans.”

Amer. Journ. Anat., Vol. vii, Part 3, 1908, pp. 409-421.

12. Lewascuew, S.—< Ueber eine eigentiimliche Veranderung der Pankreaszellen

warmbliitiger Tiere bei starker Absonderungsthatigkeit der Drie.”

Arch. f. Mikr. Anat., Bd. xxvi, 1886, pp. 453-485.

- 13. Massarr.—*Sul pancréas di pesci.’ Rend. R. Accad. dei Lincei, Vol. vii,

Fase. 5. 1898, pp. 1384-137.

14. Opr, KE. L.—‘‘ Disease of the Pancreas.” (Lippincott) Philadelphia, 1903.

“The llistology of the Islands of Langerhans.” Johns Hopkins

Bull., Baltimore, Vol. vi, p. 117.

16. Oprrrt, A—‘ Lehrbuch der Vergleich. Mikr. Anat. Wirbeltiere.” Jena, Bd. iii,

1900, pp. 800-818.

17. Renyiz, J.—‘‘The Epithelial Islets of the Pancreas in Teleostei.” Quart.

Jour. Micros. Science, Vol. xlvii, 1904-1908, p. 379.

Op. cit.,

OS Oe GSI

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43.

44.

49.

GAS GLANDS OF SOM® TELEOSTEAN FISHES. 243

References to “‘ Red Body” and other Literature.

Bagtront, 8.—* Zur Physiologie der Schwimmblase der Fische.” Zeitschr. f.

Allg. Physiologie (Verworn), Bd. viii, 1908, p. 1.

Bror.—*‘ Mémoire sur la Nature de l’Air contenu dans la Vessie Natatoire des

Poissons.” Mémoires de Physique et de Chemie de la Société

@Arcueil, T. i, 1807, Paris.

Bour, C.— The Influence of Section-of the Vagus Nerve on the Disengage-

ment of Gasesin the Air-bladder of Fishes.” Journal of Physiolosy,

Vol. xv, 1894, p. 494.

“Sur la Sécrétion de ’Oxygéne dans la Vessie Natatoire des Poissons.”

Compt. rend. Acad. Sci. Paris, T. cxiv, 1892, p. 1560.

Bripex, T. W.—“ The Structure and Function of the Air-bladder in certain

Fishes.” Proc. Birmingham Phil. Soc., Vol. vii, 1889-1891, p. 144.

Fishes, in ‘ The Cambridge Natural History,’ Vol. vii, 1904.

Brxowsk1, L. & Nusspaum, J.—* Beitriige zur Morphologie des parasitischen

Knockeutisches, Fierasfer Cuv.” Bull. Acad. Sci. Cracovie, 1904,

p. 409.

Coee@1, A.—“ Intorno ai corpi rossi della vescica natatoria di aleuni Teleostei.”

Mitt. Zool. Stat. Neapel, Bd. vii, 1886, p. 381.

— “Ueber den epithelialen Theil der soe. Blutdriisen in der Schwimm-

blase des Hechtes (Wsox lucius).” Morph. Jahrb., Bd. xy, 1889,

p. 555.

ConFIGLIAcut.—Sull’ analisi dell’ aria, contenuta nella vescica natatoria dei

Pesci.” Pavia, 1809.

Cornine, H. K.—“ Beitrige zur Kenntnis der Wundernetzbildungen in dem

Schwimmblasen der Teleostier.” Morph. Jahrb., Bd. xiv, 1888, p. 1.

Detnexa, D.—‘ Zur Frage ueber den Bau der Schwimmblase.” Zeit. wiss.

Zool., Bd. Ixxviii, 1904, p. 149.

DELAROCHE.—‘ Observations sur la vessie aérienne des Poissons.” Ann.

Mus. d’Hist. Nat., T. xiv, 1809, p. 184.

pE Seapra, A. F.—“Sur les Corps Rouges des Téléostéens (Note pré-

liminaire).” Bull. Mus. d’ Hist. Nat. Paris, T. iti, 1897, p. 217.

Drascu, O.—Archiv f. Anat. u. Phys., Physiol. Abth., 1889, p. 96. :

Emery, C.—*Le Specie del Gencre Fierasfer.” Monog. 2, Fauna u. Flora

des Golfes von Neapel, 1880.

Gourrer, E.—“Du Role de la Vessie Natatoire.’ Ann. Sci. Nat. (8),

Zoologie, T. vi, 1866.

GuyENor, H.—* Contribution 4 l'étude anatomique et physiologique de la

Vessie Natatoire des Cyprinides.” Compt. rend. Soc. Biol., T. viii,

1905, p. 794.

Hatpane, J. 8.—‘“Secretion and Absorption of Gas in the Swimming-

Bladder and Lungs.” Science Progress, Vol. vii (Vol. ii, n. s.) 1898,

pp. 120 & 237.

Howe tt, W. H.— A Vext-book of Physiology.’ Philadelphia, 1909.

Hurner, G.—‘ Zur physikalischen Chemie der Schwimmblasengase.” Arch.

f. Physiol., 1892, p. 54.

Humporpr & Provengan.—Schweigger’s Journal, 1809, Bd. 1, p. 119.

Mém. de la Soc. d’Arcueil, I’. ii, 1809, p. 400.

Humponpr.— Reise in die Aequinoctialgegenden Deutsche Bearbeitung.’

Stuttgart, Bd. 1, 1859.

Jacops, Cur.— Ueber die Schwimmblase der Fische.” Inaug.-Dissert.,

Tiibingen, 1898.

Tubinger Zool. Arb., Bd. iii.

JarGer, A.—* Die Physiologie und Morphologie der Schwimmblase der

Fische.” Pfluger’s Archiv f. Physiol., Bd. xciv, 1903, p. 65.

——— “Die Physiologie der Schwimmblase der Fische.” Biol. Centralbl.,

3d. xxiv, 1904, p. 129. -

—— “Die Schwimmblase der Fische.” Bericht der Senckenbergischen

Naturforschenden Gesellschaft in Frankfurt-am-Main, 1904 (Oct.

1903).

—— “Zur Physiologie der Schwimmblase der Fische.” Anat. Anzeig.,

Bd. xxix, 1906, p. 683.

—— “Hrwiderung aut die in Heft 7/8 dieses Bandes des Anatomischen

Anzeigers erschienene Entgegnung von Frau Reis und Herrn Nusbaum

(Krakau) : “Zur Physiologie der Schwimmblase der Fische.’” Op.

cit. Bd. xxx, 1907, p. 588.

Miiye-Epwarps, H.—‘Legons sur la Physiologie et Anatomie Comparée.’

T. 1, 1857, p. 368 e¢ seq.

59.

DR. W. N. F. WOODLAND ON THE

Morzau, A.—“‘Sur VAir de la Vessie Natatoire des Poissons.” Comot. rend.

Séances Acad. Sci., 1863.

— “Recherches expérimentales surles Fonctions de la Vessie Natatoire.”

Ann. Sci. Nat. (6), Zoologie, I’. iv, 1876.

——— ‘Mémoires de Physiologie.’ Paris, 1877, T. xv, p. 494.

Miter, J.—‘ Ueber Nebenkiemen und Wundernetze.’ Archiv f. Anat.,

Physiol. u. Med., 1840, p. 101.

Nussavum, J., & Reis, K.—“ Beitrage zur Anatomie und Physiologie des sog.

Ovals in der Schwimmblase der Fische.” Bull. Acad. Sci. Cracovie,

Sci. Math. et Nat., 1905.

Nussavum, J.—“ Zur Histologie der tatigen Gasdrtise und des Ovals bei den

Teleostiern (eine Antwort an Alfred Jaeger).’ Anat. Anzeiger,

Bd. xxxi, 1907, pp. 169-174.

Opret, A.—Article on Atmungs-Apparat: Schwimmblase, in ‘ Ergebnisse

der Anatomie und Entwickelungsgeschichte,’ Bd. xiv, 1904, edited by

Fr. Merkel und R. Bonnet, Wiesbaden.

——— Op. cit., Bd. xv, 1905.

Op. cit., Bd. xvii, 1907.

——— Article on Atmungs-Apparat: Schwimmblase, in the ‘Lehrbuch der

Vergleich. Mikr. Anat. Wirbeltiere.’ Bd. vi, 1905, pp. 104-164.

Prierer.— Ueber die Funktion der Schwimmblase.’ Berlin, 1892.

QuEKET?, J.—‘On a Peculiar Arrangement of Blood-Vessels in the Air-

bladder of Fishes, with some Kemarks on the Evidence which they

afford of the True Function of that Organ.” ‘Trans. Mier. Soc.

Lond., Vol. 1, 1844, p. 99.

Rets, K., & Nussaum, J.—* Zur Histologie der Gasdriise in der Schwimm-

blase der Knockenfische, zugleich em Beitrag zur Trophospongien-

frage.’ Anat. Anzeiger, Bd. xxvii, 1905, p. 129.

——— “Weitere Studien zur Kenntnis des Baues und der Funktion der

Gasdriise und des Ovals in der Schwimmblase der Kaiockenfische

(Ophidid, Percide).” Op. cit., Bd. xxviii, 1906, p. 177.

Rets, C.—‘ Weitere Beitrage zur Kenntnis der Gasdrtise bei den Knocken-

fischen.” Bull. Internat. Acad. Sci. Cracovie, 1906, p. 771.

—— ‘“Materyaly do mortfologii i fizyologi pecherza plawnego ryb

kostnoskieletowych.” Krakéw, Rozpr. Akad., 1906, pp. 639-670.

(“ Materialien zur Morphologie und Physiologie der Schwimmblase

der Teleostier.” Abhandl. d. Akad. d. Wissensch. Krakau, Bd. xlvi,

1907.)

Reis, K. & Nussaum, J.—“ Erwiderung auf den von Dr. A. Jaeger in Bd. 29,

No. 24 des Anatomischen Anzeigers verotfentlichten Artikel: ‘ Zur

Physiologie der Schwimmblase der Fische.” Anat. Aunzeiger,

Bd. xxx, 1907, pp. 204-207.

RicwHarD, J.—* Note sur le Gaz de la Vessie Natatoire des Poissons.” Compt.

rend. Acad. Sci. Paris, TI’. exx, 1895, p. 745.

Snore, T. W.—“ Notes on the Origin of the Liver.” Jour. Anat. Physiol.,

Vol. yv, n. s., 1891.

“On the Development of the Renal-Portals and Fate of the Posterior

Cardinal Veins in the Frog.” Op. cit., Vol. xvi, n. s., 1901.

Tuto, O.—< Die Entstehung der Schwimmblasen (Vorlautige Mittcilung).”

Biol. Centralbl., Bd. xxiii, 1903, p. 528.

— — “Die Entstehung der Schwimmblasen.” Korrespondenzbl. d. Naturf.-

Ver. Riga, Bd. xlvii, 1904, p. 26.

—— “Die Luttwege der Schwimmblasen.’’ Zool. Anzeig., Bd. xxx, 1906,

. 591.

Resa eeMiak Gaerne M.—‘ Ricerche sui gas contenuti nella vescica natatoria

dei Pesci.’ Nota 1 in; Atti Accad. Lincei Rend., Vol. ii, 1887,

2. sem.; Nota 2 in: ibéd., Vol. iv, 1. sem. ; Nota 3 in; ibid.

——— “Ueber die Gase in der Schwimmblase der Vische.” Arch. f. Physio-

logie, Jahrg. 1889, p. 54.

Vincent, S., & Barnes, A. S.—* On the Structure of the Red Glands in the

Swim-bladder of certein Fishes.” Jour. Anat. Physiol., Vol. xxx,

1896, p. 545.

Wooprann, W.—‘A Suggestion concerning the Origin and Significance of

the ‘ Renal-Portai System’ &c.” Proc. Zool. Soc. Lond., 1906.

—— “The ‘Renal-Portal System’ and Kidney Secretion.” Nature,

June 13, 1907, pp. 151, 152.

—— “On a New Gland in Teleostean Fishes.” Proc. Zool. Soc. Lond.,

1908, pp. 431-433.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 945

EXPLANATION OF PLATES II-IX.

[All figures of sections drawn with the aid of the camera lucida. In figures 4,9, 13,

14, 15, 26, 38, 46, 49, 52 and 57, which, like most of the others, have been greatly

reduced in size during the process of reproduction, the capillaries of the rete

mirabile have not been reduced to nearly the same extent as the other parts of

the figures, since, had this been done, they would have been invisible. ]

Anguilla vulgaris (P1. IL. figs. 1-8).

Fig. 1 (X18). Semi-diagrammatic transverse section through the anterior end of

gas bladder (A.B.) and pneumatic duct (P.D.). G.E., glandular

epithelium of bladder; D.H., epithelium of duct; A., artery, and V.,

vein supplying bladder.

2 (X18). Similar section, more posterior. The artery and vein have each

divided, each pair of vessels consisting of an artery and a vein.

3 (X18). Still more posteriorly the artery and vein of each pair have

subdivided soas to form a mass of intermingled arteries and veins.

4 (X18). Each mass of arteries and veins has, just before the pneumatic duct

joins the bladder, subdivided to form a bunch of minute parallel

arterial and venous capillaries—the rete mirabile bipolare geminum

(Miiller) here seen in transverse section.

5 X18). Where the duct joins the bladder, the minute capillaries of the rete

mirabile have reunited to a considerable extent, arteries with

arteries and veins with veins, to form large vessels, which then

recapillarize in order to supply the gas gland. P.D., pneumatic

duct which has now altered the character of its epithelium and

become part of the bladder, P.B.

6 (X 18). Semi-diagrammatic transverse section through the posterior region

of the bladder, P.B.

7 (X 500). Transverse section through the lining epithelium of the pneumatic

duct. Note the large capillaries and squamous epithelium

covering them.

8 (X 500). Transverse section through the glandular epithelium of the bladder

(gas gland). G.D., one of the gland ducts formed by the folding

of the epithelium.

Ophichthys (Sphegebranchus) imberbis (Pl. II. fig. 9).

Fig. 9 (X cir. 10). Semi-diagrammatic transverse section through “red body,”

where the duct (P.D.) joins the bladder (B.). R.M., the single

rete mirabile, the long axis of which is situated transversely in the

ventral bladder wall. The capillaries of the rete ave here seen in

longitudinal section.

Syngnathus acus (Pl. II. figs. 10-13; Pl. III. figs. 14-21),

Fig. 10 (X27). Semi-diagrammatie transverse section through the anterior

attachment of the bladder. A. and V., arteries and veins which

subdivide to form the rete mirabile ; Ves., vessels not taking part

in the formation of the rete ; B.G., masses of the diffuse pancreas.

11 (X 27). The arteries and veins are here subdividing to form the rete mirabile.

Figs. 12, 13 (X 27). The formation of the rete mirabile (R.M.).

Fig. 14 (X 27). The anterior end of the bladder (A.B.) has just appeared. The

rete mirabile (R.M.) is fully formed.

15 (X 27). Thearterial and venous capillaries of the rete mirabile have reunited

to form larger vessels which then recapillarize to supply the

bladder epithelium (G.E.). B., bladder cavity.

Figs. 16-18 (X 27). The vessels of the rete mirabile gradually disappear in supplying

the bladder epithelium.

Fig. 19 (X 500). A fold of the “unthickened” bladder epithelium. B.L., gland

duct between folds of epithelium; G.S., granular matter present in

only very minute quantity ; V.D., darkened zone of cytoplasm

round capillary wall. ,

20 (X cir. 1000). Transverse section through rete mirabile. Venous capillaries

indistinguishable from arterial. S.G., disintegration-products of

the erythrocytes.

21 (X 1600). Stages in disintegration of the red blood corpuscles situated in the

capillaries of the gas gland. S.G. as in fig. 20.

Proc. Zoou. Soc.—1911, No. XVII. 17

246 DR. W. N. F. WOODLAND ON THE

Gobius niger (Pl. III. fig. 22; Pl. IV. figs. 23-32, 34; Pl. V. fig. 33).

Fig. 22 (x 1). Appearance of the “ red body ” in the ventral bladder wall.

23 (X 16). Semi-diagrammatic transverse section anterior to the “red body.”

A. and V., the initial artery and vein which form the rete mirabile ;

B.G., pancreas masses ; O.L., outermost layer of the bladder wall;

G.E., glandular epithelium lining bladder cavity. It will be

noticed that the vessels and pancreas lie outside the bladder wall.

24 (X 16). More posterior section. The initial artery and vein have subdivided

to a small extent. The outermost layer of the bladder wall has

become broken through by the arteries, veins and masses of

pancreas.

25 (X 16). The arteries and veins, intermingled with masses of the pancreas,

are rapidly subdividing to form the rete mirabile and the whole

mass is now practically situated inside the bladder wall.

26 (X16). The rete mirabile (R.M.) has now commenced to supply the

glandular epithelium (G.E.) and is situated wholly inside the

bladder wall, z. e. internal to the outermost layer (O.L.). B.G.,

portion of the pancreas remaining external to the bladder.

27 (X 330). Three folds of the bladder glandular epithelium in transverse

section. C.R.M., minute capillaries of the rete mirabile; BU.,

gas-bubble; IC.C., intracellular capillary.

28 (X 800). Gas gland cell with intracellular capillary (IC.C.). Note the

striped darkened zone of cytoplasm next the capillary.

29 (X 800). Gas gland cells with inter- and intracellular capillaries (IC.C.).

30 (X 800). Gas gland cell with intracellular lumen in transverse section.

Intracellular ducts, when thus seen, are not always easy to

distinguish from bubbles.

31 (X 800). Gas gland cell with intercellular lumen (INT.C.L.) in transverse

section.

32 (X 1000). Transverse section through anterior end of rete mirabile, showing

the curious endothelium (END.) of the arteries in this region and

the large amount of erythrocyte-disintegration granular matter

contained in the veins. The arteries (A.) are here distinguishable

from the veins (V.) by their smaller size, by their thicker walls, by

possessing the curious endothelium and by being practically free

from granular matter (8.G.).

33 (X 800). A duct of the pancreas in longitudinal section. The numerous

nuclei and the syncytial character of the wall are noticeable. A

large amount of granular secretion-matter is present in the duct.

34 (x 800). A duct of the pancreas in transverse section.

Gobius minutus (Pl. V. fig. 35).

Fig. 35 (X 1000). Portion of the folded epithelium of the gas gland in a very active

condition. Bubbles are seen being produced (usually near the

nuclei) in the interiors of most of the cells, others are being

extruded from the cells and one is shown lying freely in the

bladder cavity. The largeamount of granular matter lying in the

bladder lumen external to the gas gland cells which has resulted

from the bursting of extruded bubbles is noticeable.

Siphonostoma typhle (rondeletii) (Pl. V. fig. 36).

Fig. 36 (X 1000). In addition to the erythrocytes there are shown three of the

curious “ white ” corpuscles which are so striking in appearance in

this fish, the cytoplasm being very dense. These corpuscles are

numerous.

Peristethus cataphractus (Peristedion cataphractum) (Pl. V. figs. 37-41).

Fig. 37 (X 1). Appearance of “red body” in ventral wall of bladder.

38 (X 16). Semi-diagrammatic transverse section across the anterior end of the

“yed body” (see A—B, text-fig. 57, p. 202). S.E., squamous ‘

epithelium lining the greater part of the bladder; G.E., glandular }

epithelium; R.M., rete mirabile. r

Figs. 89-41 ( 1600). Cells of the gas gland showing formation of bubbles in

cytoplasm.

GAS GLANDS OF SOME TELEOSTEAN FISHES. 247

Trigla hirundo (Pl. V. fig. 43; Pl. VI. figs. 42, 44).

Fig. 42 (x 1). Lateral “red bodies” (L.R.M.) as seen in ventral wall of bladder

(=lateral divisions of the elongated “red body,’ the median

divisions being hidden by the muscle-bands, M.B., in the middle).

G.E., the glandular epithelium (too broad in figure).

43 (X 16). Semi-diagrammatic transverse section through the middle of the

““yed body” represented in fig. 42. M.B., two strong muscle-

bands on each side of the middle line; R.M., rete mirabile ;

L.R.M., lateral rete mirabile; G.E., glandular epithelium ;

S., strand of nerve-fibres supplying muscles.

44 (xX 250). Glandular epithelium of bladder in transverse section. B.L., gland

duct.

Smaris vulgaris (Pl. V. fig. 46; Pl. VI. fig. 45).

Fig. 45 (xX 1). Appearance of “red body” in ventral wall of bladder.

46 (X 16). Semi-diagrammatic transverse section across anterior end of “red

body.” R.M., rete mirabile; G.E., gas gland epithelium; S.E.,

general squamous epithelium lining bladder; B.L., bladder cavity.

Smaris maurit (Pl. VI. fig. 47).

Fig. 47 (X 250). Pseudo-massive epithelium of gas gland in transverse section. As

explained in the text, this type of epithelium is without doubt a

derivative from the folded type and so differs from truly massive

types of gas gland epithelia which are not secondarily derived from

a folded condition. ‘The perivascular spaces seen in the figure are,

in most cases at least, due to contraction.

Ophidium barbatum L. (PI. VI. figs. 48-50).

Fig. 48 (x 1). Appearance of “red body” in ventral wall of bladder.

49 (X 16). Semi-diagrammatic transverse section across “red body.” S.E.,

squamous epithelium; G.E., gas gland epithelium; R.M., rete

mirabile. Outer layers of the bladder wall stripped off.

50 (X 800). Large gas gland cells with intracellular ducts (I1C.D.), which

possibly represent the passages of bubbles to the exterior. B.L.,

bladder cavity ; IN'T.C.D., intercellular duct.

Box boops (Pl. VI. fig. 51; Pl. VII. fig. 52).

Fig.51 (x 1). Appearance of “red body ” in ventral wall of bladder.

52 (X cir. 27). Semi-diagrammatic transverse section through the middle of the

*‘yed body” shown in preceding figure. R.M., rete mirabile;

S.E., squamous epithelium; G.E., gas gland epithelium; B.iu.,

bladder cavity.

Atherina hepsetus (Pl. VI. fig. 53; Pl. VII. figs. 54, 55).

Fig. 53 (xX 1). Appearance of “red body” in ventral wall of bladder (only the

vessels and retia mirabilia are seen).

54.(X 1000). The glandular epithelium in transverse section.

55(X 1000). The glandular epithelium of a young Atherina (15 mm. iong)

showing incipient division of the cells in a plane at right angles

to the bladder wall. C.T.C., connective tissue cells. Notice the

large size of the nuclei as compared with those in the adult gland

of another specimen. Though no capillaries are shown in the

figure, yet they are plentiful in most parts of the gland.

Coris julis (Pl. VI. fig. 56; Pl. VII. figs. 57, 58).

Fig. 56 (xX 1). Appearance of “red body ” in ventral wall of bladder.

57 (X 16). Semi-diagrammatic transverse section of “red body.” Lettering as

before.

58 (X 500). The glandular epithelium in transverse section. B.L., bladder

cavity ; S.E., superficial squamous layer of the glandular mass ;

BU., gas bubble; INT.C.D., narrow intercellular lumen.

17*

248 — ON THE GAS GLANDS OF SOME TELEOSTEAN FISHES.

Corvina nigra (Pl. VII. fig. 59).

Fig.59 (X 1). Appearance of “red body” in ventral wall of bladder.

Sargus rondeletii (Pl. VIII. fig. 60).

Fig. 60 (x 1). Appearance of “red body ” in ventral wall of bladder.

Balistes capriscus (P1. VIII. fig. 61).

ig.61 (X 1). Appearance of “red body” in ventral wall of bladder. The “ red.

body” is, in this case, situated posteriorly in the bladder, not

anteriorly.

Zeus faber (Pl. VIII. figs. 62-64; Pl. IX. fig. 65).

r.62 (X 1). Appearance of “red bodies’ in ventral wall of bladder. R.M.., rete

mirabile; G.H., gas gland; M., anterior muscular band.

63 (X 875). Transverse section across base of massive gas gland showing syn-

cytial masses which probably represent the regenerating portion of

the gland. Cell-outlines only become visible in the more peripheral

regions. The syncytial nuclei divide by ordinary mitosis and are

of several sizes. giant nuclei bemg among them.

64 (x 666). Amitotic division of nuclei in some small giant cells, intermediate-

sized cells and small cells of the gas gland. The nucleolus seems

to be the first part of the nucleus to divide.

65 (X 875). A binucleated giant cell surrounded by the ordinary small cells..

Compare the magnification of this with the last figure.

Gadus morrhua (Pl. VIII. fig. 66).

Fic. 66 (x 27). Semi-diagrammatic transverse section across “red body.” R.M.

s) “, on : . 1 . . . Bi

tufts of rete mirabile; G.H., gas gland epithelium; M.C., cap of

connective tissue; B.L., bladder cavity.

Cepola rubescens (Pl. VIII. fig. 67; Pl. IX. fig. 68).

Fig. 67 (x 1). Appearance of “red body ”’ in ventral wall of bladder.

68 (X 333). Glandular epithelimm in transverse section. The perivascular

darkening of the cytoplasm is here very marked. The ducts of

the gland are very thin and inconspicuous.

Perca flwiatilis (Pl. TX. figs. 69-72).

Fig. 69 (X 1). Appearance of branched “ved body ” in ventral wall of bladder, the:

portions of gas gland forming margins to the small fan-shaped

retia mirabilia.

70(X 1000). A small portion of the gas gland of Perca (fixed with Zenker):

showing the evolution of bubble-masses from the cytoplasm in

certain cells of the epithelium, consequent upon the experiment of

weighting the fish (Appendix B, cf. text-fig. 61, p. 240).

71 (X 1000). Cells of the gas gland of Perca (fixed with the Glacial-Absolute:

mixture) exhausted by the activity of the gland (cf. text-fig. 62).

One of the bladder-like structures is shown in connection with one

of the cells.

72 (x 1600). Unexhausted, 7. e. inactive cells of the gas gland of Perca (fixed

by the same method) for comparison with those of fig. 71.

Nerophis equorius (Pl. IX. fig. 73).

Fig. 73 (X 666). Five gas gland cells of Nerophis in contact with acapillary. This.

figure, coloured to resemble my preparations as far as possible,.

shows in the gas gland cell the striped zone of cytoplasm next the .

capillary wall (indicative of absorption) and the erythrocyte-

disintegration granular material present in the blood.

ON SKULLS OF OXEN FROM NEWSTEAD. 249

12. On Skulls of Oxen from the Roman Military Station

at Newstead, Melrose. By J. C. Ewanr, M.D., F.R.S.,

E.Z.S.*

[Received and Read February 7, 1911.]

(Text-figures 63-91.)

Professor Marcellin Boule, in his recent work on the Grotto of

Grimaldi, states that Cuvier, Riitimeyer, Nehring, Gaudry,

Boyd Dawkins, Duerst, and other naturalists who have studied

the Quaternary Bovide have regarded the Urus (os tawrus

primigenius) as identical with our modern Los taurus, of which it

was probably the ancestor, and from which it differed only by its

greater size.

Though many naturalists since the days of Cuvier have

directed their attention to the history of Domestic Cattle, the last

word has not yet been said about their origin, hence in dealing

with the remains of cattle from the Roman Military Station

at Newstead, Melrose, the investigator must still bear in mind

that a final answer has not yet been given to the question—Are

modern European cattle descended from the Urus, Bos taurus

primigenius® When discussing the origin of British cattle

Prof. Hughes remarks: “ Czsar mentions that there were large

herds of domesticated cattle in Britain, and we know from

numerous excavations into Roman and Roman-British rubbish-

pits that these belonged not to the Urus but to Los longifrons.

This, then, is the native breed with which we must start in all our

speculations as to the origin and development of British oxen.

The Romans found that breed here and no other.”

Writing about the Celtic Shorthorn (Los longifrons Owen,

Bos brachyceros Riitimeyer) Mr. Lydekker says, ‘‘ It is, and can be,

nothing but a variety of Bos tawrus” derived from the wild Urus

at a very remote epoch—“ the occurrence of remains of an

apparently similar breed in the prehistoric lake-dwellings of

Switzerland suggests that the breed may have been established

prior to the separation of Britain from the Continent” ¢.

Bos frontosus Nilsson, Lydekker also regards as a variety of

the Urus, and as there was no other primitive Wild Ox in Europe,

and an Eastern derivation being in the highest degree improbable,

Lydekker says that all the domesticated breeds of European

cattle must trace their ultimate ancestry to Bos primigenius.

While satisfied that the Domestic Cattle of Europe are descended

from Bos primigenius, Lydekker thinks it is quite probable that

the origin of the humped cattle of India (Los indicus) may be, at

least in part, different.

* The author is indebted to Mr. James Curle, Priorwood, Melrose, for the use of

the Oxen skulls found at Newstead.

+ ‘Les Grottes de Grimaldi, tome i. fascicule iii., 1910.

+ ‘ Wild Oxen, Sheep, and Goats,’ p. 18, 1898.

Text-fig. 63.

250 PROF. J. C. EWART ON

Though Professor Fairfield Osborn thinks that the Domestic

Ox, instead of being a direct descendant of Bos primigenius, is a

descendant of Bos trochoceros of the Italian Pleistocene, he differs

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in no essential point from Lydekker, and, like Lydekker, believes

British Shorthorn Cattle are descended from an indigenous

Occidental race domesticated in Europe by the Neoliths*. <A

somewhat different view is taken by Prof. Boyd Dawkins, who

* Osborn, ‘ The Age of Mammals,’ 1910.

From a skull in

The horns occupy nearly the same position as in Antelopes.

maxillze (J2.) and nasals (WVa.).

the Royal Scottish Museum,

SKULLS OF OXEN FROM NEWSTEAD. 251

Text-fig. 64.

Front part of the skull of a Buffalo (Bos bubalus).

The premaxillz (P.Z.), as in the Anoa, extend upwards between the maxillze-

and nasals.

Text-fig. 65.

Front part of skull of the Urus (Bos primigenius).

In this skull the premaxille are only slightly connected with the nasals, but in a

skull in the British Museum the premaxille extend nearly as far upwards

between the maxillze and nasals as in the Buffalo (text-fig. 64).

2D) PROF. J. C. EWART ON

says: “It is clear the domestic animals were not domesticated in °

Europe, but that they had already been under the care of Man

probably for long ages in some other region. ‘The Turf-Hog, the

Celtic Shorthorn, the Sheep, and the Goat have been domesticated

in the countries in which their wild ancestors were captured by

the hunter in Central Asia. To this region also belong the

Jackal, the Wild Boar, and the Wild Horse, and in ancient times

the Urus. It is therefore probable that all these domestic animals

came into Europe with their masters from the south-east—from

the Central plateau of Asia—the ancient home of all the present

European peoples.”

Text-fig. 66,

Skull of a Urus in the Anatomical Museum, University of Edinburgh.

The premaxilla has only a slight connection with the nasal, the forehead is flat, the

temporal fossa is closed behind, and the horn-cores curve forwards at nearly

a right angle to the forehead.

Of Continental zoologists, Prof. Duerst has for some years been

directing most attention to the origin of domestic animals. In

his report on the Animal Remains found in Turkestan by the

SKULLS OF OXEN FROM NEWSTEAD. 253

Pumpelly Expedition, Duerst says: “‘ My recent studies on fossil

remains of the bovines of the Indian Pleistocene have shown

Text-fig. 67.

Front part of skull of American Bison.

The premaxillz are short and far removed from the nasals.

From a skull in the Anatomical Museum, University of Edinburgh.

Text-fig. 68.

illite

Nil

Front and side views of the skull of a young foetal Ox. Nat. size.

The premaxille: (P.I/.) reach the nasals (Wa.), the frontals (#%.) extend nearly as

far backwards as in the adult Anoa, and there is a large gap between the

frontals (#7.), parietals (Pa.), and supra-occipital (Oc.).

254 ON SKULLS OF OXEN FROM NEWSTEAD.

me that the Indian (Narbada and Siwaliks) and China Taurina

are the exact equivalents of the European Urus (Bos primigenius

Bojanus), excepting some very slight variations produced by

different geographical and local influences, so that the Bos nama-

dicus Falconer and Cautley would represent the Kuropean Urus

for the Asiatic Continent, especially the North Indian mountains

and their neighbourhood” *.

In recapitulating the results of his studies of the bovids of

Anau, Duerst says:—‘ In the lower layers of period Ia from

— 24 feet upwards there occur the remains of a wild Bos namadicus

Falconer and Cautley. During period Id there originates from

this wild form a domesticated bovid, large and stately, provided

with long horns. Judging from the measurements of the

preserved bones this is absolutely the same Ox that was possessed

by the Ancient Egyptians.

“In the period II the size of the animal seems to have some-

what diminished, unless possibly a smaller bovid reached Anau

with the other newly imported domestic animals. It is, however,

possible that this form of cattle of the culture IT originated in

adecline of the cattle-breeding of the later Anau-li; as, indeed,

the originally large long-horned Ox of the early Babylonians had

already become small and short-horned in Assyrian times, and

to-day, after a relatively shorter interval, shows a tendency to

become hornless ” 7.

Duerst goes on to say :—“ The first remains of the long-horned

breed (Bos tawrus macroceros) belong at Anau about 8000B.c. We

find the same animal again about 3000 to 4000 B.c. in Babylonia

and Egypt. At about 6000 B.c., however, we find that the large

long-horned animal of Anau has become small and small-boned

and has developed into a short-horned breed (Bos brachyceros) t.

Therefore all who do not believe in an autochthonous domesti-

cation of the animals for each separate culture-sphere must admit

that the original large and stately long-horned Ox of Anau was

spread by tribal migrations before 6000 B.c. to Persia and

Mesopotamia and into Egypt and Central Africa on the one hand

and on the other to India and Hastern Asia, where according

to Chinese accounts it arrived in 3468 B.c.

“ Did the migration of the West occur only after the small

breed had become established, 2. e. about 6000 B.c., or even between

6000 and 7000 B.c. when the turbary sheep had formed? To this

question we have as yet no answer. We must, however, add that

it was not in Anau alone that through unfavourable conditions

of life the originally large and stately Ox was changed into the

stunted and short-horned form (Bos taurus brachyceros). The

same change took place in Mesopotamia, as one may easily per-

ceive in comparing the long-horned cattle of Chaldean or Sumero-

Accadian times with the Assyrian small short-horned and the

* “Animal Remains from Excavations at Anau,’ Carnegie Institution of Wash-

ington, p. 361.

+ Duerst, op. cit. p. 369.

{ Bos brachyceros is the same as Bos longifrons.

Text-fig. 69.

Oc.

-A.—Front view of a fcetal Ox skull about the fifth month. The premaxille (P.I.)

reach the nasals .(a.), the frontals (Fr.) are long, the parietals (Pa.)

narrow, and the interparietals (Ip.) lie between the parietals and supra-

occipital. Nat. size. J

‘B.—F ront part of same skull, to show relation of premaxillz to nasals. Nat. size.

256 PROF. J. C. EWART ON

modern loose and short-horned or hornless cattle. There is,

therefore, no reason for rejecting the assumption or hypothesis

that the Ox of Anau, which about 7000 3B.c. was undergoing

this change of form, finally reached Central Europe, after its

migration through Southern Russia and Eastern Europe, in the

stunted form of Los taurus brachyceros,” * 7. e. in the small Celtic

Shorthorn generally known in England as Bos longifrons.

If the conclusions arrived at by Duerst and others are justified,

it follows (1) that all the modern domestic cattle—the humped

breeds of India and Africa as well as the European breeds—are

derived either from the Urus of Europe (os taurus primigenius)

or its reputed near relative the Urus of Asia (Los namadicus), or

are a blend of varieties or races of these two species; and (2) that

modern British breeds have been formed by crossing the Celtic

Shorthorn (Los longifrons vel brachyceros)—the small domesti-

cated race widely distributed over Britain in pre-Roman times—

with Continental breeds (including short-horned as well as long-

horned varieties) introduced since the Roman invasion. Are these

conclusions supported by the remains of cattle from the border-

fort occupied by Roman auxiliaries during the first and second

centuries of the present era ?

Prof. Boule, in his recent work on the Grimaldi fossils, says the

genus os (which includes the most specialized members of the

Bovid family) seems to have been represented in Pliocene times

by Los planifrons and Bos acutifrons of the Siwalik deposits of

India. Duerst regards Los planifrons t as the ancestor of both

Bos primigenius aa Bos namadicus, bat Rutimey er thinks that

though Los planifrons may be ancestral to, it is only a variety of

Bos primigenius, the Kuropean variety of Los namadicus.

Though Los primigenius, like the Bison, only reached Europe

in Quaternary times, it was soon widely distributed—its

remains occur in English Pleistocene deposits containing Hlephas

antiquus and in deposits of a like age over the greater part of

Europe and also in North Africa. While Gos primigenius was

extending its range over Europe, Los namadicus was spreading

over Asia.

Tt has hitherto been supposed that the Bison was more abun-

dant in Europe during Pleistocene times than the Urus, but

Prof. Boule believes that, at least in the vicinity of Grimaldi, the

Urus was from the first as common as the Bison.

About the colour of the Urus nothing absolutely certain is

known, but from drawings of Medizval, as well as Paleolithic

artists we can forma fairly accurate conception of its conformation,

A picture, believed to have been made in Bavaria about 1500 A.p.,

probably brings out the chief points of Bos prinigenius t.

* Duerst, op. cit. p. 440.

+ Accor ding to Duerst’s latest view there is no real difference between Bos plani-

frons of Riitimeyer and Bos acutifrons of Lydekker, but at one time he believed

Bos acutifrons was the predecessor of Bos namadicus, to which the Bibovine (Gaur

and Banting) group and especially the Indian Zebu were related.

+ This picture is reproduced in the ‘ Cyclopedia of American Agriculture,’ vol. iii.

1900 ; the Urus apparently survived in Poland up to 1627.

SKULLS OF OXEN FROM NEWSTEAD. 257

Text-fig. 70.

} |

| if

DM |

A.—Front view of Calf’s skull at birth.

The premaxillze (P.JL) fail to reach the nasals (Na.). The parietals and inter-

parietals have coalesced to form a narrow plate (Pa.) between the frontals

(Fr.) and supra-occipital.

B.—Front part of same skull.

The premaxilla (P.W.) is short and some distance from the nasal (Wa.).

Text-figs. 69 & 70, from specimens belonging to Prof. Charnock Bradley, D.Sc.

258 PROF, J. C. EWART ON

The Urus was widely distributed in Britain in Neolithic times,

but the examination of Roman and Roman-British stations has

hitherto afforded no evidence that Bos primigenius still survived

in England at the Roman invasion.

A number of more or less complete skulls of the Urus have

been found in Seotland, and fragments of skulls, limb-bones, or

horn-cores believed to belong to the Urus have been found

in almost every county in Scotland between the Solway and the

Pentland Firths, and some horn-cores found in Orkney are so

large that it is assumed they belong to Bos primigenius.

Text-fig. 71.

Front part of skull of a polled Aberdeen-Angus Ox, with premaxillee (P. I.) extending

nearly as far up between the maxillee (1/~.) and nasals (Wa.) as in the Buffalo

(text-fig. 64, p. 251). From a specimen in the Royal Scottish Museum.

Though there is no evidence that the Urus survived long

enough in England to give rise to the Chillingham and other

“‘ wild” white park cattle, it has been suggested that a sufficient

number survived in the Caledonian forests to found the Cadzow,

Atholl, or other Scottish herds of ‘ wild” cattle. The bones from

Newstead, however, afford no evidence that the Urus still survived

in Scotland when the Romans constructed the border-fort during

the later part of the first century A.D.

The skull of Los taurus primigenius is in some respects more

highly specialized than that of any other member of the Bovide.

Hitherto in studying bovine skulls a very considerable amount of

attention has usually been directed to the position, size, and

SKULLS OF OXEN FROM NEWSTEAD. 259

direction of the horns. I shall, however, especially refer to the

premaxille, occiput, and temporal fossze.

1. The Premazxille.—tIn some Zebras (e. g. the true Burchell

Zebra) the premaxilla may only be connected with the nasal for

a distance of 12 to 15 mm., while in others (e. g. a variety which

lives near Lake Baringo) the connection between the nasal and

the premaxilla may exceed 50 mm. Differences in the food may

account for the premaxille being long in some cases and short in

others. The premaxille may require to be firmly wedged in

between the maxillz and nasals in varieties in the habit of feeding

during part of the year on coarse hard food, but only slightly

connected with the nasals in varieties which usuaily consume soft

green herbage.

Text-fig. 72.

Skull of a Syrian Ox with vestigial horn-cores and a forehead like that of flat-polled

Aberdeen-Angus cattle; the premaxillze (P..) as in text-fig. 71 extend far

up between the maxillee (Mzx.) and nasals (Na.). From Wilckens, ‘ Natur-

geschichte der Haustiere.’

In the Anoa (Los depressicornis) of Celebes and in the wild

Indian Buffalo (Los bubalus) the premaxille are firmly wedged in

between the maxille and nasals. In the Anoa (text-fig. 63, p. 250)

the total length of the premaxilla is 105 mm.,and its connection with

the nasal is 35 mm.; in the Buffalo (text-fig. 64) the total length

of the premaxilla may be 173 mm., and its connection with the

nasal 40 mm.

This long intrusion of the premaxilla between the nasal and

260 PROF. J. C. EWART ON

maxillary bones doubtless increases the strength of the front

part of the jaw which supports the horny pad against which the

lower incisors bite. As the Anoa is partial to the neighbourhood

of water, it probably feeds on coarse grasses like its ally the

Tamarau of the island of Mindoro, which is said to browse on

sugar-cane.

The Indian Buffalo lives in the neighbourhood of swamps and

jungles, and probably also feeds on reeds and coarse grasses—

food which necessitates long, firmly secured premaxille.

In the Catalogue of the Ungulata in the British Museum

Dr. Gray says that in the true Oxen “the premaxille are large

and always extend upwards into the triangular space between the

maxille and the nasals and consequently articulate with both

these bonesasin Lubalus ; in the genus Sibos, on the other hand,

the premaxille are small and are attached only to the distal

extremity of the maxilla and are separated by a considerable

interval from the nasals.” As will appear below, the premaxillee,

instead of being always long enough in true Oxen to extend

upwards between the maxille and nasals as in the Buffalo, are

sometimes so short that they fail to reach the nasals as in the

Bison (text-fig. 67, p. 253).

In Bos primigenius the premaxille, in all the skulls I have

examined, reach the nasals. In some cases the connection with

the nasals is only 5 mm., but in a Urus skull in the British

Museum the premaxille extend nearly as far upwards between

the nasals and the maxille as in the Buffalo (text-fig. 64). The

extent of the connection between the premaxilla and the nasal

in a Urus skull in the Anatomical Museum of the University of

Edinburgh is shown in text-figs. 65 & 66.

The premaxilla is shorter and further removed from the nasal

in the Bison than in any other member of the Bos genus (text-

fig.67). According to Major Heber Percy, the European Bison

‘Care fond of grazing on a coarse aromatic kind of grass known as

Zubr grass.” Others state that ‘“ they are equally fond of browsing

on the leaves, young shoots, bark, and twigs of trees,” and that

“in winter they are driven to subsist entirely on buds, twigs, bark,

and such patches of dry grass and fern as remain” *,

The prairie Bison of America apparently fed chiefly on grass,

hence doubtless the necessity for the extensive migrations, but the

American woodland Bison seem to ‘‘ subsist chiefly on the leaves

and twigs of the birch and willow”. It is conceivable that

owing to the softer nature of the food of the Bison the necessity of

having the premaxille firmly wedged in between the nasal and

maxillary bones no longer exists.

In a very young Domestic Ox skull (breed unknown) the pre-

maxilla (text-fig. 68) bears the same relation to the nasal as in

the Urus represented in text-fig. 66; in an older skull it occupies

* Lydekker, ‘ Wild Oxen, Sheep, and Cattle,’ p. 77.

+ Lydekker, op. cit. p. 91.

SKULLS OF OXEN FROM NEWSTEAD. 261

the angle between the maxilla and the nasal (text-fig. 69, A, B);

but im a skull believed to be that of a new-born calf there is

a considerable gap between the premaxilla and the nasal (text-

fig. 70, A,B). In some of the skulls from the Newstead Fort the

premaxille are large and have nearly as extensive a connection

with the nasals as in the Buftalo; in others they are short and

separated from the nasals by a considerable interval.

'Pext-fig. ay,

Na.

[LEY

Front part of skull of polled Newstead Ox in which the upper end of the

premaxillae (P.1.) bifurcates.

In a Newstead polled skull the premaxille are long and more

extensively connected with the nasals than in the Urus skull

shown in text-fig. 66. In some Aberdeen-Angus cattle the pre-

maxille are short, in others they are so long and so extensively

connected with the nasals that they reach to within 25 mm. of

the lachrymal bones (text-fig. 71); in a Buffalo with premaxille

of the same length (text-fig. 64) the Jachrymals are 75 mm. from

the premaxille. Apparently in some Syrian cattle the premaxille

extend well up between the maxille and nasals. Ina skull of a

Syrian Ox figured by Duerst* the premaxille bear the same

relation to the nasals and lachrymals as in the Angus skull

figured. As this Syrian Ox had small imperfect horns (text-

fig. 72), and as the skull apart from the horn-cores resembles

the skull of the flat-polled Angus strain, it may represent a phase

in the evolution of the Aberdeen-Angus breed. The large

* Wilckens, ‘ Naturgeschichte der Haustiere,’ p. 311, 1905.

Proc. Zoo. Soc.—1911, No. XVIII. 18

262 PROF. J. C. EWART ON

premaxille, coupled with the fact that the parietal extends

well on to the forehead, suggest that Aberdeen-Angus cattle

have in part sprung from a race domesticated in Central Asia.

While in two of the skulls of black polled Aberdeenshire cattle

hitherto examined the premaxille have an extensive connection

with the nasals, in the skull of a white polled “ wild” Cadzow Ox

as in a black polled Galloway, the premaxillz bear practically the

same relation to the nasals and the lachrymals as in Los prim-

genius, represented in text-fig. 65.

Text-fig. 74.

Front part of a horned Newstead skull in which the premanxillze (P.JZ.) are short

and fail to reach the nasals (Va.).

In most of the Newstead skulls examined the premaxille are

absent, but when the maxille are present it is usually possible to

say whether or not the premaxille had reached, or all but reached,

the nasals. They probably reached the nasals in at least 90 per

cent. of the skulls of the Celtic Shorthorn (Los longifrons) type,

and in about 70 per cent. of the skulls belonging to long-horn and

cross-bred animals. In the Celtic Shorthorn skulls the premaxille,

though long and almost in contact with the nasals, instead of

ending in a wedge-shaped process, are bifurcated and send a

process backwards over the maxilla as wellas one upwards between

the maxilla and the nasal (text-fig. 73). In having the proximal

end forked the Celtic Shorthorn skulls agree with the skull of an

SKULLS OF OXEN FROM NEWSTEAD. 263

Ox from Irkutsk figured by Duerst*. In cross-bred cattle the

premaxilla may be forked on the one side, as in text-fig. 73, but

long and rounded on the other as in text-fig. 71.

Text-fig. 75.

Front part of skull of a horned Cadzow Ox with notched premaxille (P.IZ.) which

fail to reach the nasals (Wa.). From a skull in the Royal Scottish Museum.

Text-fig. 76.

Front part of skull of cross-bred Shorthorn in which the premaxillie (P.Z.)

fail to reach the nasals (WVa.).

As already mentioned, Dr. Gray stated in the British Museum

* Wilckens, ‘ Naturgeschichte der Haustiere,’ p. 305, 1905.

18*

264 PROF. J. CG. EWARL ON

Catalogue of the Ungulata that in the true Oxen ‘“ the premaxillee

are large and always extend upwards between the maxille and

nasals and consequently articulate with both these bones as in

Bubalus.” lLydekker, in discussing the premaxille in the Bovide,

points out that in Bos chinensis, a new species described by Gray

in 1870 (t.e. before the publication of the Catalogue of the

Ungulates), ‘the premaxille are small and do not extend upwards

to the nasals precisely as in Libos”*. But while in Bos nama-

dicus and in Los chinensis the premaxille may be always small, they

are not invariably small in the Bibovine group. In a Gaur skull

in the Royal Scottish Museum the premaxille reach the nasals,

and Mr. Pocock informs me that, of four Gaur skulls he examined

Text-fig. 77.

Skull of polled Newstead Ox with forked premaxille, large orbits, deeply notched

occiput, uneven forehead ending in a rounded mesial prominence. Skulls of

this type with horns are sometimes said to belong to Bos frontosus Nilsson.

in the British Museum, one has the premaxille in contact with

the nasals, while in another skull they are separated by a long

interval from the nasals. Of all Lydekker’s statements about the

premaxillee, the one which bears most on the present enquiry is to

the effect that a small premaxilla is “ never found in any Huropean

Ox” +. If the premaxilla is smallin Los namadicus, but never

small in European cattle, the presumption is that Kuropean cattle

are not descended from Los namadicus, but from Los primigenius,

in which the premaxille in all the skulls examined reach the

nasals.

In one of the Newstead skulls, which probably belongs to an im-

ported Ox, as large as a modern Aberdeen-Angus steer, the total

length of the premaxilla i is only 118 mm. —47mi. shorter than the

* Memoirs of the Geological Survey of India, series x. vol. 1. p. 19.

+ Op. cit. p. 3.

265

SKULLS OF OXEN FROM NEWSTEAD.

text-fig.

The distance between the prema

premaxilla of the Aberdeen-Angus heifer represented

this Newstead Ox is 80 mm., in the Aberdeen-An

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5 mm.

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cus only

xilla and the lachrymal

to reach the nasal and makes no attempt to fill up the gap between

But in addition to being small, or rather because it is small, it fails

(al

"81 SY-9X9],

266 PROF. J. C. EWART ON

the maxilla and the nasal (text-fig. 74). As it happens, there is

a skull of one of the so-called “ wild” white Cadzow cattle from

Hamilton Park in the Royal Scottish Museum which probably

belonged to an Ox including the Celtic Shorthorn amongst its an-

cestors. In this Ox (text-fig. 75), though the premaxilla is 142mm.

in length it neither reaches the nasal nor yet extends into the gap

Text-fig. 79.

Fr.

Oc. Crest

F.M.

FM.

Outlines of the occiput and part of the forehead of three Urus skulls.

Fr., frontal; FIL, upper border of foramen magnum.

In all three skulls the intercornual ridge projects far beyond occipital condyles.

From skulls in the Anatomical, Antiquarian, and Free Church College

Museums, Edinburgh.

between the maxilla and nasal. In shape and in being notched at

its proximal end, it reminds one of the premaxilla of the Celtic

Shorthorn (text-fig. 73). Another skull with premaxillee which fail

to reach the nasals may be mentioned, because it has many of the

SKULLS OF OXEN FROM NEWSTEAD. 267

characteristics of the Urus. In this skull (text-fig. 76), which

belonged to a cross-bred Shorthorn, the premaxilla in shape closely

agrees with the premaxilla of the Urus, butit is relatively shorter

and neither reaches the nasal nor extends into the space between

the nasal and the maxilla,

Text-fig. 80.

Front view of skull of Bison (Bos bison).

Tn the Bison the horn-cores are near the orbits, the parietals (Pa.) are not overlapped

by the frontals (Fr.), and the premaxillze (P..) are far removed from the

nasals (Wa.).

Though very few skulls have been available for study, it is

evident (1) that in some modern European cattle as well as in some

of the cattle in Britain during the Roman occupation, the pre-

maxille were short and failed to reach the nasals, and (2) that as

268 PROF. J. C. EWART ON

the premaxille vary in wild as well as in domestic cattle the relation

of the premaxill to the nasals is of little diagnostic value.

The Forehead.—In the Newstead skulls the forehead varies

considerably ; in some specimens it is flattened as ina typical Urus

(text-fig. 66), and the ridge between the horn-cores is nearly

straight; in others there are prominences and depressions and the

intercornual ridge is arcuated as in some specimens of the Urus,

while in a polled skull (text-fig. 77) of the frontosus type, the vertex

projects forwards and upwards to form a well-marked rounded

mesial prominence. In this polled skull there is also a projection

from the middle of the forehead and a well-marked ridge at each

Text-fig. 81.

Hind part of skull of the Urns represented in text-fig. 66.

The front of the horn-core is in a line with the occipital condyle, and the temporal

fossais closed behind by a plate of bone which supports the horn-core.

side between the orbit and the long deep orbital suleus. In old

animals the parietals seem to be completely covered by the

frontals. But even when the intercornual ridge projects fax

beyond the level of the occipital condyles the interparietal

probably enters into the formation of the forehead. That the

parietals (or the interparietals) form part of the forehead in polled

Oxen is indicated by text-fig. 78 B, Pa. That in horned Oxen the

parietals also enter into the formation of the forehead is suggested

by a large skull of the Urus type in the Royal College of Surgeons

Museum, London (No. 1121 A). In this skull the frontals are

separated by a wedge-shaped piece of bone (probably the inter-

SKULLS OF OXEN FROM NEWSTEAD. 269

parietal), which forms the middle portion of the intercornual

ridge. Oscar Schmidt * states that ina front view the parietals

“cannot be seen at all,” but probably in all cases the parietals

in the Ox, as in the Bison (text-fig. 80), enter into the formation

of the forehead. The great specialization of the frontal region of

the skull in the true Oxen is made evident by the examination of

foetal skulls. In avery young foetus the frontals (text-fig. 68, /’r.)

only form the front portion of the roof of the cranium; but about

Text-fig. 82.

Frontlet and horn-cores of a small Newstead Ox with a prominent forehead.

The horn-cores curve upwards and backwards.

the fifth month they are relatively very much larger than the

parietals (text-fig. 69, /’r.), and at birth they form nearly the entire

roof of the cranium (text-fig. 70). After birth the frontals

continue to grow backwards, and they eventually form all but the

wedge-shaped middle portion of the intercornual ridge. In the

Urus the frontals sometimes project 100 mm. (4 inches) beyond

the occipital condyles (text-fig. 79), but in the Aberdeen-Angus

the occiput almost forms a right angle with the frontals, and the

* “The Mammalia,’ p. 176.

Text-fig. 83.

270 PROF. J. C. EWART ON

parietals (text-fig. 78 B), as in the Bison (text-fig. 80), obviously

enter into the formation of the forehead. The highly speciaiized

condition of the occiput in the Urus has resulted from the horns

having migrated upwards from their original position above the

qi i i \

\y\\! i)

Occiput of Urus skull represented in text-figs. 66, 79 (outline showing occipital crest, Oc. Cvrest.), and 81.

orbits(text-fig 63) to lie entirely behind a line passing through the

occipital condyles (text-fig. 81). In addition to Newstead skulls

with the forehead flat or presenting ridges or prominences, there

is a skull (text-fig. 82) in which the forehead is so convex that the

shallow; C1, occipital condyle; Z.N., shield for attachment of ligamentum nuche; I.R., intercornual ridge. -

The occiput is quadrangular, the occipital crest flattened, and the notches (V.) between the crest and horn-cores are

The occiput is completely cut off from the temporal fossa.

Text-fig, 84,

SKULLS OF OXEN FROM NEWSTEAD. 271

horn-cores at their origin ave far behind the most prominent part

of the forehead. Moreover, the horn-cores in this case are flat-

tened and curve outwards and backwards as in some Eastern

eattle. In their position and direction the horns in this Newstead

Ox decidedly differ from the horns (also of the flat type) of a

Shetland heifer in my possession, which project almost directly

upwards as in some Zebus.

\ AS \ \

Wie

"UY?

74,

The occipital crest is rounded, and deep notches (V.) separate the supra-occipital from the horn-cores.

€

Occiput of the Newstead skull represented in text-fig. 86, p.

The horn-cores ave, as a rule, circular in section and, in skulls of

the Celtic Shorthorn type, curve outwards and forwards nearly

at aright angle to the frontals. In other skulls they curve out-

wards and upwards, or outwards and downwards, or outwards,

occipital crest; O., occipital condyle; P.P., paroccipital process ; Z.N., surface for attachment

A deep semilunar depression occupies the middle of the intercornual ridge immediately above the

of ligamentum nuche ; I.R., mesial frontal prominence.

ibe, PROF. J. C. EWART ON

upwards, and backwards as in some Indian cattle. Except when

they curve forwards at right angles to the frontals, as in typical

Celtic Shorthorns, the horns assist but little im settling the race

to which the Newstead skulls belong.

The Occiput.—In some Newstead skulls the occiput has the

characteristics of the Urus represented in text-fig. 83. The general

outline is quadrangular, the occipital erest is flattened but not

encroached on by the temporal fossze, the lateral borders are nearly

straight, and a line carried through the highest point of the crest

lies below the centre of the horn-cores; further, there is a shield-

like projection (Z..V.) for the insertion of the ligamentum nuche.

The distance from the crest to the lower border of the foramen

magnum is nearly the same as the distance between the condyles

(C.) and little more than half the distance between the notches

(V.Y.) below the horn-cores on a level with the temporal fossie.

- Text-fig. 85.

sei EONS Ee

Occiput and horn-cores of a Urus in which the notches below the horn-cores

are deeper than in text-fig. 83.

This occiput closely resembles the occiput of a Bos namadicus skull

in the British Museum.

The nearly smooth supracristal part of the occiput extending

between the horn-cores measures from above downwards about

half as much as the part lying between the crest and the lower

border of the foramen magnum. In some cases the supracristal

part is flat and nearly in a line with the occiput proper, in others

it projects beyond the crest (text-fig. 79) and, though concave

in the centre immediately over the shield for the ligamentum

nuche, it is prominent and convex above and forms a marked

angle with the forehead. In some cases the upper border of the

SKULLS OF OXEN FROM NEWSTEAD. 273

occiput viewed from behind is nearly straight, in others it is

slightly concave or distinctly arcuated.

In specimens ot the Urus with very thick horn-cores a bridge of

bone extends downwards from the base of the horn-core and forms

a posterior wall for the temporal fossa (text-figs.81 & 83, V.), which

completely cuts off the fossa from the occiput. It is owing to the

temporal fossee being closed in behind by extra supports for the

horn-cores that the width of the occiput immediately below

the horn-cores is sometimes nearly twice as much as the distance

between the crest and the lower border of the foramen magnum.

Though in the majority of the Newstead skulls the occiput

conforms to the Urus type there are several with short premaxille

in which the occiput reminds one of Lydekker’s Los acutifrons.

Writing in 1880 Lydekker pointed out that Bos acutifrons of the

Punjab Siwaliks had no marked relationship to any existing species

and was ‘ widely different in the form of its occiput, frontals, and

horn-cores from Bos primigenius” *.

Though Riutimeyer and others regarded Los planifrons and Bos

acutifrons as varieties of a species closely allied to Bos primigenius,

no skulls of true Oxen have hitherto been described in which the

occiput resembles that of Los acutifrons.

In Bos namadicus, as well asin Los primigenius, the notches

below the bases of the horn-cores are shallow, but in the Newstead

skull represented in text-fig. 84 indentations or notches (V.)

below the horn-cores are so deep that the connection between the

upper (parieto-frontal) part of the occiput and the lower (infra-

cristal) part is relatively short. Because of these indentations the

occiput viewed from behind (text-fig. 84) bears a resemblance to

that of the Gaur (Bos gaurus).

In Bos primigenius the summit of the occipital crest is on a

lower level than the middle of the horn-cores (text-fig. 83),

but in the Newstead skull of the acutifrons type a line carried

through the summit of the rounded arch formed by the occipital

crest (text-fig, 84) passes above the middle of the horn-cores.

At the widest part the occiput of the first century skull

(text-fig. 84) measures 215 mm., and from the lower border of

the foramen magnum to the occipital crest the distance is

115mm. The depth (115) multiplied by 100 and divided by

the width (215) gives an index of 53:5.. In the Urus represented

in text-fig. 85 the corresponding index is 47. Between the

notches under the horn-cores the distance in the Newstead

skull (text-fig. 84) is 123 mm.: 123x100+215 (the total

width of occiput) gives an index of 57; in the Urus, owing to the

shallowness of the notches under the horn-cores, the correspond-

ing index may be 90. The distance (115 mm.) between the

lower border of the foramen magnum and the occipital crest

multiplied by 100 and divided by the distance (123 mm.) between

the infracornual notches gives an index of 93:5; the correspond-

ing index in the Urus may be only 52:4.

* Memoirs of the Geological Survey of India, series x. vol. i. p. 2.

274 PROF. J. C. BWART ON

Text-fig. 86.

A.—Front view of the Newstead skull of which the occiput is represented in

text-fig. 84, p. 271.

B.—Temporal fossa of above skull. The fossa communicates freely with the

occiput. Compare with fossa of the Urus (text-fig. 81, p. 268).

SKULLS OF OXEN FROM NEWSUEAD. 275

Text-fig. 87.

A.—Front view of a white “ wild”? Cadzow Ox from Hamilton Park.

B.—Occiput and horn-cores of the above skull.

This skull closely resembles the Newstead skull (text-figs. 84 & 86) with a deeply

notched occiput. From a specimen in the Royal Scottish Museum.

276 PROF. J. GC. EWART ON

From the narrow shield-shaped projection (..V.) under the

evest (text-fig. 84), a narrow ridge extending towards the foramen

magnum divides the lower part of the occiput into two equal

portions roughened for the attachment of muscles. Between the

crest and the upper border of the foramen magnum the occiput

is distinctly concave—near the crest ip is excavated to a depth

of 12mm. The part of the occiput external to the condyles (C.)

is divided by a ridge continuous with the incurved paroccipital

processes (P.P.) into two irregular, more or less concave, rough

surfaces. The greatest width across the occipital condyles (C.),

is 99 mw. (only 23 mm. less than the space between the

infracornual notches): 99x 100+123 (the width of the occiput

under the horn-cores) gives an index of 80. The corresponding

index in the Urus skull (text-fig. 83) is only 55.

Text-fig. 88.

Occiput end horn-cores of an American Dison.

The frontals neither cover the parictals (text-fig. 80, p. 267) nor project beyond the

level of the occipital crest, and the horn-cores are separated by wide gaps

from the supra-occipital.

The part of the occiput above the occipital crest is divided into

three portions, viz.:—a mesial semicircular deeply excavated

portion and two lateral portions continnous with the horn-cores

externally and the forehead above. The mesial semicircular

concave portion (text-fig. 84), 60 mm. wide and 20 mm. deep, has

a rough margin, 3 to 6 mm. in thickness, which forms the nearly

horizontal border of the mesial frontal protuberance (text-

fig. 86, A).

From the measurements given it follows that the occiput of

SKULLS OF OXEN FROM NEWSTEAD. Dit

the Newstead skull (text-fig. 84) differs profoundly from that of

the more highly specialized Urus skull represented in text-fig. 85.

The Horn-cores.—The horn-cores of the Newstead skull,

characterized by a deeply notched occiput, are pyriform as in

Bos acutifrons—they measure at their origin 60 mm. from above

downwards and 45 mm. from before backwards. At a distance

of 30 mm. from the beginning of the grooved surface they

measure 56 mm. by 42 mm. The fragments of the horn-cores

present (text-fig. 86, A) are convex above, grooved in front and

below, and extend outwards and slightly forwards.

The Temporal Fossa.—In the Urus skull figured the temporal

fossa is completely closed behind (text-fig. 81) by a wall of bone

which helps to support the horn-core (text-fig., 83); the external

opening of the fossa, rounded and contracted behind, is pointed

in front and arched above (text-fig. 81). In the Newstead skull

with short premaxille and a notched occiput, the temporal fossa

is wide behind, where it opens on to the occiput, wider than in

the Urus in front, and the upper border is sinuous (text-fig. 86, B)

as in the Bison.

The Base of the Craniwm.—In the Urus the anterior as well as

the posterior tubercles are well developed, but in the Newstead

skull (text-fig. 84), as in Bos acutifrons, the anterior tubercles are

small and inconspicuous.

In having the lower part of the occiput separated from the

upper by deep notches under the horn-cores the Newstead skull

(text-fig. 84) decidedly differs from bos primigenius (text-fig. 83)

and also, though to a less extent, from Gos namadicus. Again, in

having the occipital crest overhanging the true occiput, the

Newstead skull differs from the Gaur and Banting.

Tf Lydekker is right in assuming that the occipital crest in

Bos acutifrons “ extends upwards to within a short distance of

the vertex craniw so that the supra-cristal portion of the occipital

region is reduced to a very narrow band” *,the Newstead skull

also differs from os acutifrons. If, however, in Bos acutifrons

the mesial depression immediately below the vertex is supra-

cristal, the Newstead skull with a deeply notched occiput may be

regarded as belonging to a race allied to, or descended from, Bos

acutifrons of the Punjab Siwaliks.

It will doubtless be asked, Ave any of the modern breeds of

cattle characterized by short premaxille and an occiput of the

primitive type represented in text-fig. 84? As it happens, the

skull (text-fig. 87) of the white “ wild” Cadzow Ox with short

premaxille (text-fig. 75) has an occiput (text-fig. 87, B) of the

acutifrons type, and otherwise resembles several of the Newstead

skulls. The skull of the Cadzow Ox seen from behind looks

almost as if it were intermediate between the Bison (text-fig. 88)

and a Urus with fairly deep notches (text-fig. 85).

At one time the Hamilton Park herd of white cattle consisted

* Memoirs of the Geological Survey of India, ser. x. vel. i. p. 27.

Proc. Zoou. Soc.—1911, No. XIX. 19

278 PROF. J. C. EWART ON

almost entirely of polled individuals, now they are said to be all

horned *. There is a tradition that Cadzow cattle re-acquired

horns through a Highland bull, which, for a time, took forcible

possession of the herd, and it is a matter of history that some

years ago a “wild” Chillingham bull was made use of to rein-

vigorate the Cadzow “ wild” cattle. The intercrossing, perhaps,

led to reversion towards a race in the possession of, if not

actually introduced by, the Roman auxiliaries who garrisoned the

border-fort during the later part of the first century 7.

Text-fig. 89.

Front part of the skull of a Zebu (Bos indicus) in which the premaxillie

reach the nasals.

In addition to Newstead skulls with an occiput of the Urus

and acutifrons types, there are skulls which in the occiput

agree with certain Indian cattle. In Indian as in European

domestic cattle the premaxille are sometimes long and in

contact with the nasals (text-fig. 89), sometimes short and

terminating some distance from the nasals (text-fig. 90). In

at least some Zebus long premaxille are correlated with a wide

Urus-like occiput, and short premaxille with a narrow deep

occiput. In several small Newstead skulls the occiput closely

* A skull of a polled Cadzow Ox in the Anatomical Museum of the University

of Edinburgh agrees in the premaxille and occiput with Bos prinigenius.

+ The Chillingham and Chartley “ wild” park cattle in their occiput conform to

the Urus type.

SKULLS OF OXEN FROM NEWSTEAD. 279

agrees with a Zebu skull (text-fig. 91) in the Royal College of

Surgeons Museum, London. In Bos primigenius the shield-like

projection for the Ligamentum nuche is in contact with the

occipital crest, but in some of the small Newstead skulls, as in

the skull of the acutifrons type, the rough surface for the attach-

ment of the ligamentum nuche lies, as in some Zebus, nearly

midway between the occipital crest and the upper border of the

foramen magnum. Further inquiry may show that Indian

domestic cattle are in part descended from ancestors allied to

Lydekker’s Bos acutifrons and that the Celtic Shorthorn (Bos

longifrons Owen, Bos brachyceros Riitimeyer) is intimately related

to some of the small Oriental races.

Text-fig. 90,

Front part of the skull of a Zebu in which the premaxillze fail to reach

the nasals.

The Origin of the Newstead Oxen.

In addition to cross-bred animals Newstead has yielded five

fairly distinct types of Oxen, viz.:—(1) Oxen of the Celtic Short-

horn type; (2) Long-horned Oxen of the Urus type; (3) Oxen

with an occiput of the Los acutifrons type; (4) Oxen with a

convex forehead, an arcuated intercornual ridge and horns curving

outwards and backwards; and (5) Hornless Oxen. It is im-

possible to say definitely how any of these forms originated, but

it may be safely assumed that they were not all formed in

Europe from Los primigenius.

Sys

280 PROF. J. C. EWART ON

Though Bos primigenius reached Kurope from Central Asia in

early Pleistocene times, there is no evidence that it was living

under domestication on the arrival in Post-Pleistocene times of

the Neoliths. Further, there is no evidence of the existence of a

small Ox of the Celtic Shorthorn type in European Paleolithic

deposits—Prof. Boule ¢. g., found not a single fragment of Owen's

Bos longifrons in deposits of the Reindeer age at Monaco.

Neither is there any evidence of the existence of a small wild Ox

in Pleistocene times in Central Asia.

Text-fig. 91.

Occiput of the Zebu with the premaxillx reaching the nasals (text-fig. 89).

In several of the Celtic Shorthorn skulls from Newstead the occiput resembles

that of the Zebu.

Text-figs. 89-91 from skulls in the Royal College of Surgeons Museum, London.

The examination of the bones of Oxen from Anau, Turkestan,

led Duerst to conclude :—(1) that a large long-horned breed was

formed by the Anau-l about 8000 B.c. from a large wild Asiatic

vace which he regarded as the exact equivalent of the Kuropean

Urus (Bos primigenius) ; (2) that about 6000 B.c. a small short-

horned breed, identical with Owen’s Los longifrons and Riiti-

meyer’s Bos brachyceros, was formed at Anau, or brought to Anau

from some other settlement in Central Asia. If, as seems

probable, the Urus was the only wild Ox in Central Asia in

prehistoric times, it must be assumed that the small Ox in the

SKULLS OF OXEN FROM NEWSTEAD. 28]

possession of the Anau-li was a dwarfed descendant of an Asiatic

variety of Bos primigenius *.

It is doubtless possible that a small breed may have been

formed out of the huge Urus by the Neoliths immediately after

they reached Europe, but the evidence, so far as it goes, suggests

that the Neoliths brought the “Celtic” Shorthorn with them

from Central Asia.

Though in Britain the Urus was hunted by the Neoliths—

evidence of this we have in the Urus skull from Burwell Fen,

near Cambridge, with the frontals pierced by a Neolithic flint

implement—there is no evidence that Bos primigenius was once

domesticated in Britain or that the Neoliths allowed their

domestic cattle to breed with young wild bulls.

But on the Continent the Urus was apparently domesticated at

a comparatively early period and crossed with the small breed

originally brought from Central Asia. Hence it may be said

that up to at least the Bronze age the majority of the domestic

cattle in Europe were the descendants of Bos primigeniws—some

being nearly pure descendants of the imported ‘ Celtic” Short-

horn breed, while others were pure or nearly pure descendants

of the indigenous wild Urus (Bos taurus primigenius).

There is no evidence that there existed in Europe or in Central

Asia a variety of Bos primigenius with the occiput deeply notched

and otherwise resembling the one represented in text-fig. 84,

Neither is there any evidence that in Bos namadicus—the Urus

of India—the occiput was deeply notched or characterized by an

excavated intercornual ridge. The only extinct form to which

the Newstead skull represented in text-figs. 84 and 86 bears any

marked resemblance is Bos acutifrons of the Pliocene Siwaliks.

Tt has been suggested that the Newstead skull with a deep mesial

semicircular depression above the occipital crest belonged to a

hybrid between an Ox and a Bison, but this view is not supported

by the skulls of Ox-Bison hybrids. Moreover, in its occiput and

premaxille this Newstead skull (text-figs. 84 & 86) very closely

agrees with the skull of a Cadzow Ox (text-fig. 87) in the Royal

Scottish Museum. It may hence in the meantime be assumed

that some of the cattle in the south of Scotland during the

Roman occupation were descended from an Indian race allied to

Bos acutifrons.

Of the Newstead cattle with horns curving backwards and down-

wards (text-fig. 82) it need only be said that they seem to be more

intimately related to Bos namadicus than to Bos primigenius.

The polled Newstead cattle represent two distinct types. Some

had a nearly flat forehead, a nearly straight “intercornual ” ridge,

and a square-shaped occiput; in others the forehead was very

uneven and ended in a pronounced mesial prominence which

projected upwards and forwards (text-fig. 77). The Newstead

* The small Ox of Anau is probably now represented in Asia by the long-browed

Zebus characterized by small horns of the Bos longifrons type.

282 MR. G. P. FARRAN ON COPEPODA

cattle with a flat poll (as in the modern Galloway and in poiled

Cadzow cattle) obviously belong to the Urus type, while those

with a mesial prominence seem to belong to the frontosus type of

Nilsson. As there were hornless cattle in Egypt as early as the

Fourth Dynasty, we are not likely soon to ascertain where or

when polled breeds originated. There is no evidence that any

of the varieties of polled cattle are descended from hornless wild

ancestors; neither is there certain evidence of a pure horned

race suddenly producing polled offspring. If, as seems highly

probable, the absence of horns is not due to reversion, the polled

condition was either acquired by the gradual reduction in the size

of the horns or suddenly, 7. e. by mutation. In polled breeds

which now and then produce individuals with “ loose horns ” the

polled condition was perhaps acquired slowly, while in polled

breeds in which “ scurrs ” are unknown the polled condition was

perhaps due to a mutation. Though the Galloway and the white

polled “wild” Cadzow cattle may have descended from Sos

prinigenius, the white polled Somerford, and the round polled

Aberdeen-Angus cattle with the premaxille extending well up

between the maxille and nasals, may be the descendants of an

Oriental race allied to a modern Syrian breed apparently in the

act of losing the horns.

13. Plankton from Christmas Island, Indian Ocean.—I. On

Copepoda of the Family Coryceide. By Grorer P.

FARRAN™.

[ Received October 31, 1910: Read February 7, 1911. |

(Plates X.—XTV. 7)

Through the kindness of Dr. W.T. Calman I have had the oppor-

tunity of examining the Copepoda of a small collection (8 bottles)

of Plankton, made by Sir John Murray, K.C.B., F.R.S., and

Dr. C. W. Andrews, F.R.S., at Christmas Island in the Indian

Ocean, and presented to the British Museum by Sir J. Murray.

The gatherings were all made at approximately the same time

(July—August, 1908) and in the same locality, on the north side

of the island in shallow water near shore, and for this reason

it has not been thought necessary to refer to each of them

separately.

The collection, though small in bulk, is exceedingly rich in

species, and the genus Corycwus is especially well represented.

It has been recognized that there are, in the the genus Cory-

ceeus, two groups differing from each other in several distinct

characters, the most notable being the form of the ventral

process, situated between the maxillipedes and the first pair of

* Communicated by Dr. W. T. Canman, F.Z.8.

+ For explanation of the Plates see p. 296.

Paes SLoly ce

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FROM CHRISTMAS ISLAND. 283

feet of the female, which is beak-shaped in one group and semi-

circular in the other. There can be no doubt that these characters

are, collectively, of generic importance, and I propose the name

Corycella for that subdivision, as typified by C. gibbulus, which

is marked by the possession of a beak-shaped ventral process,

leaving the name Coryceus for the remaining species, of which

CO. speciosus may be selected as the type.

The genus Coryceus was founded by Dana in 1845 (6), his

diagnosis being repeated in 1846 (7), but no species belonging to

the genus was described till 1849 (8). Dana’s definition of the

genus includes both the genera defined above, and his papers

describing the species, first published without figures in 1849 (8),

and afterwards with figures in 1852 (9), contain forms belonging

to both Corycella and Coryceus, but do not indicate any one of

them as forming the type of the genus.

The principal differences between Coryceus and Corycella may

be summarized as follows :—

Genus CoRYCELLA, nov.

Abdomen of ¢ and 2 one-jointed ;

4th thoracic segment without lateral

points, not distinct from 3rd thoracic

segment. Ventral process beak-shaped

in 9. 2nd antenna with setz on 1st and

Qnd basal joints distantly feathered ;

terminal spine short in @; inner edge of

2nd basal coarsely feathered.

Ist to 3rd feet, exopodite with 0.0.1

outer edge sete.

4th foot without endopodite, exopo-

dite with 0.0.1 outer edge set.

Genus Corycaus Dana.

Abdomen of ¢ and 9 usually two-

jointed ; 3rd and 4th thoracic segments

distinctly separate, the latter with

lateral points. Ventral process semi-

circular in 2. 2nd antenna with the

setze on Ist and 2nd basal joints not

feathered ; terminal spine elongated in

&, inner edge of 2nd basal usually with

one or more strong teeth.

1st to 8rd feet, exopodite with 1.1.3

outer edge sete.

4th foot with endopodite bearing one

or two set, exopodite with 0.1.1 outer

edge sete.

The one-jointed abdomen and the feathered setze of the second

antenne occur, aS Dahl has pointed out, as characters of im-

maturity in the genus Coryceus.

Giesbrecht (10) recognises five species of Coryewus in the

group for which the name Corycella is proposed, viz., C. gibbulus

Giesbr., C. concinnus Dana, C. rostratus Claus, C. carinatus

Giesbr., and C. longicaudis Dana.’ In Dahl’s revision of the

genus (5) he accepts C. gibbulus, C. concinnus, C. carinatus,

C'. rostratus, and, under the new name of C. tenwicauda, C. longi-

caudis, asserting that Dana’s description of C. longicaudis really

refers to C. speciosus juv., a view which is strongly supported

by Dana’s figures. Dahl also recognises Dana’s (8) description of

C’. gracilis as being good, but does not give any reason for this

opinion. Dana’s description and figures of C. gracilis evidently

refer to a male of Corycella, but the specific differences between

the males of this genus are so slight that it does not appear

possible to refer the description and figures with certainty to any

particular species. Dahl’s belief that C. pellucidus of Dana is the

female of what he recognises as (’. gracilis also appears to be

a mere conjecture, there being very little evidence for or against

284 MR. G. P, FARRAN ON COPEPODA

the view in Dana’s figures, which, apparently, include two

different species.

Wolfenden (11) also believes that he has recognised the

C. pellucidus of Dana, but his figures and descriptions refer

undoubtedly to the species which Giesbrecht has described as

C. gibbulus.

Dahl, in the paper referred to above (5), which is a prelimi-

nary note on the Copepoda of the Plankton Expedition, has

drawn up a useful diagnostic table of the members of the genus

Corycceeus as recognised by. him. The portion which refers to the

genus Corycella has been tr anslated by Dr. Wolfenden (11) in his

account of the Maldive Copepoda. In this table Dahl gives as

a mark to distinguish the other members from C. rostratus,

““Viertes Beinpaar mit einem inneren Zapfen welcher eine Borste

trigt.” If by “ Zapfen” is meant endopodite, as is undoubtedly

the case in the latter part of the table, and as Wolfenden trans-

lates it, this statement is an error, as the endopodite is absent

throughout the group. There is a mistranslation in Wolfenden’s

table which detracts somewhat from its usefulness; the sentence

“Genital openings of 2 set at end of abdomen” should read

“distant from end of abdomen,” “vom Ende des Abdomens

entfernt.”

The specific characters in the genus Corycella are very slight

and depend mainly on the shape of the abdomen and the third

thoracic segment. The following is an attempt to draw up

a diagnostic table, to include the females of which a recognisable

description exists and two new species from the Christmas

Island collection, which are described below.

1. Furca only twice as long as broad...............00000ccecceeeeeeeeeee CC. vostrata.

2. Furea three or more times as long as s broad.

A. Furca shorter than widest part of abdomen in dorsal view.

a, Widest part of abdomen behind the middle, 3rd thoracic

segment with dorsal hump.

a. Abdomen less than twice as long as high im lateral

view, uneehiona fureal spine short and stout ....... C. gibbula.

B. Abdomen ca. 23: times as long as high in lateral view,

median fur i spine slender.

. Widest part of abdomen in fr ont of middle, “3rd thoracic

segment not humped dorsally.

a. Thoracic spines long, abdomen with ventral setose

pad . C. carinata.

B. Thoracic spines ‘short, abdomen without setose pad . C. curta, sp. n.

B. Furea longer than widest ‘part of abdomen.

a. Furea less than half as long as abdomen, 8rd thoracic

’, brevis, sp. n.

segment with dorsal hump ........ C. longicaudis.

b. Furea less than half as long : as s abdomen, 3rd thoracic

segment without dorsal hump Loddihaek vs Syarastene enum mang (la CONCIIIITA

CoRYCELLA GIBBULA Giesbr.

Coryceus gibbulus Giesbrecht (10).

Coryceus pellucidus Wolfenden (11).

2 Coryceus megalops Will.-Suhm (1).

Common in all the gatherings, upwards of 120 specimens.

The most easily recognised characters of the female of this

FROM CHRISTMAS ISLAND. 985

species are the deep cephalon with eyes rather small and set far

apart, the abdomen widest at its posterior third (without furca)

and usually bearing two spermatophores, the furea contained

21 times in the rest of the abdomen, the middle furcal seta thick

and equal in length to the width of the furea, and the setose pad

on the antero-ventral part of the abdomen.

Wolfenden (11. p. 1027) has described at some length a female

Coryceeus which occurred plentifully in the Maldive collections,

and which is apparently identical with the above. He ascribes

it to C. pellucidus Dana, and records C. gibbulus as a separate

species. He remarks, however, that he believes that the C. pel-

lucidus of the Maldive collections is probably identical with

C, gibbulus. His figure of the fourth foot (fig. 9, pl. xcix.) is

evidently incorrect in showing an endopodite bearing a long seta,

as in the text it is said that no endopodite is present.

Giesbrecht’s (10) figure of the female abdomen of C. gibbulus in

lateral view is not so deep nor so irregular in outline as are all

my specimens ; in this they agree more nearly with Wolfenden’s

figure, but I think there can be no doubt that all these forms are

identical.

It seems not improbable that Willemoes-Suhm’s C. megalops

was in reality this species, but his description and figure, as

given by Brady (1), can hardly be regarded as constituting a valid

diagnosis. Brady’s C. pellucidus may also include it with others.

Distribution. Red Sea, Arabian Sea, Indian Ocean (many

records), and Tropical Pacific.

CoRYCELLA BREVIS, sp.n. (PI. X. figs. 1-6, Pl. XI. fig. 7.)

Female (Pl. X. figs. 1, 2)—Length :85 mm.; cephalothorax,

in middle line, 54 mm.; abdomen and furca °3 mm.; cephalon

rather deep, curved dorsally in lateral view ; eyes large, pro-

minent in dorsal view.

2nd thoracic segment with slight dorsal hump, as in C. gibbula

and ('. longicaudis,

3rd thoracie segment (Pl. X. fig. 6) with comparatively short

lateral points.

Abdomen widest at its posterior 2? excluding furea; in lateral

view the abdomen is parallel-sided for about 2 of its length, and

is then tapered to the furca; it has a slightly spinulose antero-

ventral boss.

Furea about 6 x 1, contained 2} times in rest of abdomen ; its

median terminal spine not very strong, about twice as long as

width of furea.

Ist and 2nd antenne (Pl. XI. fig. 7) as in C. gibbula.

lst to 3rd feet (Pl. X. figs. 3, 4) almost the same as in C’. gibbula.

The 3rd joints of the exopodites of the Ist and 2nd feet are

finely serrate, in the 3rd foot the serrations are not visible.

4th foot (Pl. X. fig. 5) as in C. gibbula, hollowed on distal

margin of 2nd basal; inner margin of the 2nd basal forms an

acute angle with its base.

2986 MR, G. P, FARRAN ON COPEPODA

The characters which distinguish this species from C. gibbula

are the proportionately greater length and different form of the

abdomen and the shorter 3rd thoracic spines.

Only one specimen was found.

CORYCELLA CONCINNA Dana.

In five gatherings, about 30 specimens.

Giesbrecht (10) has given a figure of the female of this species

by which it can easily be recognised, but he does not refer to the

presence of a setose pad, as in C. gibbula, on the antero-ventral

part of the abdomen.

Distribution. Widely distributed in the Indian Ocean, Tropical

and South Pacific.

CoRYCELLA CARINATA Giesbr. (PI. XI. fig. 10.)

In five gatherings, 55 specimens.

Length of females *85 mm.

The characteristic features of this species are the cephalon (deep

from front to back), the small eyes moderately far apart, the long

slender thoracic spines, and the abdomen, widest at its anterior

fourth (excluding furea), with a ventral setose pad. The furea is

about half as long as the rest of the abdomen.

CorycELLA curRTA, sp.n. (Pl. X. figs, 7-11, Pl. XI. figs. 1-6.)

Female (Pl. XI. figs. 1, 2)—Length ‘7 mm. Very close in

general appearance to C’. carinata, but may be distinguished by

the more slender build in lateral view, by the shorter thoracic

spines, which do not reach as far as the genital openings, and by

the absence of the patch of fine setze or spinules on the antero-

ventral part of the abdomen.

The furea is short, about 2x7, and contained 24 times in the

length of the rest of the abdomen (Pl. XI. fig. 3), which is

broadest at its anterior fourth and tapered posteriorly.

The appendages have no special features. The fine serrulation,

found on the third joints of the exopods in some species, is absent

(Pl. X. figs. 7,8). The 2nd basal of the fourth foot (Pl. X. fig. 11)

is rounded on its inner face, and does not form one straight line

with the inner face of the 1st basal. The angle between the

base and inner margin of the 2nd basal is slightly obtuse.

Only one specimen was found.

In the same gathering there occurred two specimens which

seem to be the males of the above, on account of the similarity of

their thoracic spines, fourth feet and furca, and their small size.

Male (Pl. XI. figs. 4-6). Length -65 mm. Cephalothorax in

middle line ‘4 mm., abdomen and furea :°24 mm. The form of

the abdomen (Pl. XI. fig. 4) is more easily shown in the figure

than described. The outer edges of the 3rd joints of the exo-

podites of Ist and 3rd feet are not serrulate. The angle between

the base and inner margin of the 2nd basal of the 4th foot

(Pl. XI. fig. 5) is slightly obtuse.

FROM CHRISTMAS ISLAND. 287

CoRYCELLA sp., ¢. (Pl. XI. figs. 8, 9.)

There were present numerous specimens of males of Corycella

belonging to at least two species judging by size alone, as in other

characters they seemed to be in agreement. These doubtless

represent the males of C. gibbula, C. concinna, and C. carinata,

but I have not been able to refer them to their respective females.

In size the larger measured ‘85 mm. and the smaller ‘78-8 mm.

Dahl’s (5) recognition of Dana’s C. gracilis is based on a supposed

identification of a figure of a male; but Dana’s (9) figures would

apply equally well to any one of these specimens, and I do not

think that there are any grounds for regarding his description

as valid.

The proportional length of abdomen and furca in these speci-

mens was 7:3; in one of Dana’s figures of C. gracilis it is 3: 2 and

in the other 4:2. Cleve (2) has given figures of the male of

C. gibbula, which agree fairly well with my specimens, except

that the width of the abdomen is greater in his figures. He

states that the proportion of abdomen to furca is 2:1, but in one

figure shows it as 9:5, and in the other as 8:3, so it is possible

that he may have been dealing with more than one species.

The figures (Pl. XI. figs. 8, 9) are taken from a specimen of the

larger form.

Genus Coryc2us,

Of the genus Coryceus, as restricted above, there is a

considerable number of more or less well described forms, which

may be conveniently summarised under the grouping adopted

by Dahl.

Of the forms with a very long furca, a convenient but hetero-

géneous group, Giesbrecht has figured C. longistylis and C. furcifer,

while C’, lautus of Dana is easily recognised from his figure.

The species of which the females have a one-jointed abdomen

are, according to Giesbrecht, C. alatus, C. flaccus, and C. elongatus ;

to them Dahl adds C. limbatus of Brady, originally described

from a male.

Coryceeus robustus stands alone in having the end claw of the

2nd antenna elongated in both sexes.

Of the forms without sete on the genital segment of the

female, all of comparatively large size, Giesbrecht has dealt with

C. speciosus, C. dane, and C. ovalis, while Dahl accepts as a

valid description the C. vitrews of Dana.

The remaining species, which do not fall into any of the above

groups, are mostly of very small size. They all have setz at the

genital pores of the female abdomen, and may be divided into

two groups according as the endopodite of the fourth foot bears

one or two sete. Of the first group C. obtusus, C. gracilicaudatus,

and C. venustus have been figured by Giesbrecht; C. minutus,

C. catus, and OC. pacificus have been described, in his synoptic

table, without figures, by Dahl, who also admits C. latws of Dana

I88 MR. G. P. FARRAN ON COPEPODA

and (. hualeyi of Lubbock. Of the second group, with two

setze on the endopodite of the fourth foot, C. lubbocki and

C'. tenuis have been figured by Giesbrecht; C. anglicus is a well-

known species in British waters; C. amazonicus, C'. asiaticus,

C. minimus, and C. africanus figure as new species in Dahl’s table ;

and C, erythreus has been described and figured by Cleve (4).

The three new species described below belong to the last group.

Corycmus LoncistyLis Dana.

C’. longistylis Dana (9).

C. varius Brady (1).

O. longistylis Giesbrecht (10).

Present in four gatherings, 5 females and 7 males.

Both sexes of C. longistylis are easily distinguished from

C. lautus and C. furcifer, which also have a long furca, by the

broad spreading 3rd thoracic segments. The female is not com-

pletely figured by Giesbrecht, but Brady’s figure of the whole

animal (1) (pl. hi. fig. 1), under the name C. varius, is fairly

accurate.

Distribution. Indian Ocean, China Sea, and Tropical Pacific.

Corycaus LAuTuS Dana.

C. lautus Dana (6).

C. lautus Dahl (5).

Present in four gatherings, 12 females and 29 males.

The female is distinguishable from that of C. furcifer by its

larger size, 2°75 mm., C. furcifer 2 measuring only 1°38 mm.,

and by the form of the abdomen, the anal segment being much

wider in front than behind, while in C. furcifer it is only slightly

tapered. The male of C. lautws (2°15 mm.) is considerably larger

than that of C. fureifer (1:3-1:45 mm.), but otherwise resembles

it rather closely in general appearance. The furea, however, is

only 11 times as long as the rest of the abdomen, instead of

almost 14 times as in C. fureifer 3.

Both species can be readily distinguished by dissection, C. lawtas

having two set on the endopodite of the fourth foot, while in”

C. furcifer there is only one. Dana’s (9) figure of C. dautus Q is

easily recognisable as distinct from C. furcifer.

Distribution. Kingsmill I. (Dana), N.E. Atlantic.

Corycaus ALATUS Giesbrecht.

The female of C. alatus is well represented in the collection,

36 specimens occurring in five gatherings. The shape of the

abdomen, as well figured by Giesbrecht, makes it unmistakable.

The males of this species and C. flaccus present some difficulties.

While the females of C. alatus and (. flaccus number respectively

36 and 12, there are 76 specimens of a male which corresponds

closely to Giesbrecht’s (10) description of C’. elongatus 3. These

specimens are undoubtedly the males of C. alatws and C. flaccus,

FROM CHRISTMAS ISLAND. 289

but I have been unable to distinguish between them or to

separate them satisfactorily from C. elongatus ¢. They can be

readily picked out under a dissecting microscope, owing to the

fact that, apart from differences of form, the maxillipedes are

coloured a rusty-red.

Distribution. Tropical Pacific.

Corycaius rLAccus Giesbrecht.

There is nothing to add to Giesbrecht’s (10) figures and de-

scription of this species, which is easily separated from C’, alatus,

the only species which it at all resembles, by the form of the

abdomen and furea. There occurred twelve specimens, females,

in three gatherings.

The other two species of this group, C. elongates and C. lim-

batus, which Dahl (5) regards as distinct, were not found in the

collection.

Distribution. Mediterranean, Indian Ocean, Tropical Pacific.

Corycaus roBustus Giesbrecht.

C. venustus Brady (1).

C. robustus Giesbrecht (10).

Present in three gatherings, 10 males and 2 females.

The female is recognisable by its very large size and by the

form of the genital segment, which is provided with lateral ridges

and overlaps the anal segment in dorsal view.

There can be no doubt that the species which Brady (1) re-

corded as C. venusius Dana was in reality C. robustus. The

detailed figure of the abdomen shows the genital segment over-

lapping the anal, in a way which is characteristic of that species

alone. The figure (pl. liv. fig. 8) of the whole animal is

somewhat misleading, as the artist or engraver evidently, after

the picture was finished, mistook the lateral abdominal ridges

for ege-sacs, and touched them up with a view to increasing the

resemblance. Anyone who compares Giesbrecht’s figure of

C. robustus 2 (11) (pl. li. fig. 38) with Brady’s figure of C. ven-

ustus (1) (pl. liv. fig. 8) can hardly avoid the conclusion that

both had the same species before them. The dimensions given

by each are identical. Brady’s description does not mention any

characters of specific value. Giesbrecht’s suggestion that Brady's

CO. venustus was really C. obtusuws seems to me to be quite baseless.

Distribution. Red Sea, Arabian Sea, Indian Ocean, Tropical

Pacific, and off Cape of Good Hope.

Corycaus spectosus Dana.

An easily recognised species, well represented in the collection.

Tt occurred in six gatherings, 22 females and 33 imales.

Distribution. Mediterranean, Red Sea, Arabian Sea, Indian

Ocean, Tropical Pacific, off Cape of Good Hope, Tropical and

North Atlantic.

290 MR. G. P. FARRAN ON COPEPODA

Coryc&US DAN Giesbrecht.

Dahl (5) has proposed that Giesbrecht’s name of C. dance

should give way to Dana’s C. crassiusculus which he believes to

represent the male of the same species. There is some resem-

blance in the figure given by Dana to C. dane ¢, but there does

not seem to be any reason for upsetting a well-established name

with a recognisable description on account of its possible identity

with a very imperfect description with insufficient figures, to

which, at best, it can only be brought home by a process of

exclusion.

C. dane occurred in seven gatherings, 44 females and 70 males.

Distribution. Mediterranean Sea, Red Sea, Arabian Sea, Indian

Ocean, Tropical Pacific, off Cape of Good Hope.

CoRYCHUS GRACILICAUDATUS Giesbrecht. (Pl. XI. figs. 11, 12.)

Dahl (5) has proposed that Giesbrecht’s name should be re-

garded as a synonym of Dana’s C. agilis, which represents a male,

length ;!, inch or ‘83 mm. Females, which are almost. certainly

Giesbrecht’s C. gracilicaudatus, occurred in five gatherings, nine

specimens. In Dahl’s key they fall in with C. agihs.

What I believe to be the male of the species occurred in three

gatherings, 50 specimens (Pl. XI. figs. 11, 12). The length was

‘76 mm. The proportions of the abdomen and furea agreed

fairly well with Dana’s figure of C. agilis, but the 3rd thoracic

segment had the distance between its points less than the width

of the hinder end of the 2nd thoracic segment. In Dana’s figure

the poimts are spread widely outwar rds. ‘These males were

remarkable in having the longest furcal seta bright red.

Distribution. Red Sea, Arabian Sea, Indian Ocean, Tropical

Pacific.

Corycaus catus Dahl. (Pl. XII. figs. 1-3.)

Dahl (5) has given the above name to a species of Coryceus,

the female of which is only to be distinguished, according to his

diagnostic table, from C. obtusus (C. ovalis of Dahl) by having

outstanding sharp points on the ends of the 4th thoracic segment,

whereas in C. obtusus the ends of that segment are short and

blunt. The other characters, extracted from his table, are—

Endopodite of 4th foot with one seta; abdomen with seta behind

the genital opening; size not more than 1 mm.; furea compara-

tively short ; third thoracic segment spreading, more than 7 as

broad as first, the points not reaching to the distal end of the

genital segment; anal segment very little longer than its basal

width.

There are several specimens of a female Coryceus in the

collection which agree with the above diagnosis and are either

C’. catus or a new species, the probability being in favour of the

former view. They occur in five gatherings, 90 specimens in

all. The 3rd thoracic segment in dorsal view, though spreading,

FROM CHRISTMAS ISLAND. 291

is not so wide as in C. obtusus, being at its widest part, a little

behind the anterior margin, only just 3 as wide as the widest part

of the cephalon (PI. XII. fig. 1). The points of the segment are

slightly contracted, the distance between them being only 2 = of the

width of the cephalon. The total length is -9 mm.

The most noticeable features, in comparison with C. obtusus,

are the sharp slender points to both the 3rd and 4th thoracic

segments (Pl. XIT. fig. 2), and the somewhat larger eyes, separated

by rather less than their own diameter. The form of the

abdomen and furca is very similar in both species.

Corycaus optusus Dana. (Pl. XII. figs, 4-6.)

Coryceus obtusus Dana.

C. obtusus Giesbrecht.

C’. ovalis Dahl.

According to Dahl (5), Giesbrecht is in error in his identification

of the species which he has described and figured under the

name of C. obtwsus Dana, Dana’s species being different and not

found subsequently. He believes that Giesbrecht’s description

really refers to the species which Claus (2) had described under

the name of C. ovalis.

Under these circumstances the course least liable to cause

confusion would be to reject both Dana’s and Claus’s names ;

however, for the present, I assume that Giesbrecht’s name is

correct. Eighteen specimens, females, of the species in question

occurred in the collection, in three gatherings.

The distinguishing characters are the size, 1:05-1'1 mm., the

wide spreading third thoracic segments, the blunt tips to the

fourth thoracic segments (PI. XII. fig. 5), the fine setee in front

of the genital openings, the anal segment as long as it is broad

at the base and nearly as long as the furea, and the single

seta on the endopodite of the fourth foot (P]. XII. fig. 4).

Distribution. Red Sea, Arabian Sea, Indian Ocean, Tropical

Pacific, Tropical Atlantic.

Corycoaus TENUIS Giesbrecht. (PI. XII. figs. 8, 9.)

Of the forms with two sete on the endopodite of the fourth

foot and a moderately long furca there are two species, females,

in the collection. One of these is described below as new, the

other appears to be C. tenuis. In Dahl’s (5) key the choice lies

between C. anglicus and C. tenuis, the former being distinguished

in the key by the presence of a corner on the inner side of the

lateral prolongation of the 3rd thoracic segment and by a small

hook-like ventral process on the proximal end of the abdomen.

Both these characters are present in my specimens and, if the

table were closely followed, they ought to be C. anglicus. They

are, however, not that species, being distinguished from it by the

shorter genital segment with much more slender setze, the form of

the anal segment, which is parallel-sided and about. twice as long

292 MR. G. P. FARRAN ON COPEPODA

as wide, and the furea, which is slightly longer and more slender.

They agree well, on the other hand, as far as the text and figures

go, with Giesbrecht’s description of C. tenuis, though that writer

makes no mention of the ventral hook on the genital segment.

The length, 1:05 mm., is slightly greater than that given by

Giesbrecht, viz.°9 mm. The length of the cephalothorax in the

middle line, dorsal view, is ‘64 mm., of the abdomen and furca

‘41 mm., the proportional lengths of abdominal segments and

furca being 14:8:18. Measured along the ventral margin the

proportions are more nearly 12:10:17. The pigment of the

eye-apparatus shows very distinctly through the body as of a blue-

green colour.

The second antenna (Pl. XII. fig. 9) bears a long spine on the

first basal joint and a much shorter spine, one-third as long, on

the second basal joint. The inner margin of the second basal ends

distally in two strong teeth. The terminal claw of the second

antenna is just equal to the spine of the second basal.

The swimming-feet have no distinctive features, except that the

terminal spine of the exopodite of the second foot is very slightly

curved.

Six specimens were found in two gatherings.

Distribution. Indian Ocean and Tropical Pacific.

CorycHUSs DUBIUS, sp. n. (PI. XII. fig. 7, Pl. XIV. figs. 5-9.)

In the Maldive collections Dr. Wolfenden (11) obtained a

specimen of Corycceus, which, following Dahl’s table, ought to be

C. amazonicus, but, as he remarks, without figures it is impossible

to refer it with certainty to that species. In the Christmas

Island collection there occurred one specimen of a female,

described below under the name of C. dubius, which may well be

the same species as that to which Dr. Wolfenden refers, but

again the absence of figures precludes certainty. The particulars

given by Dr. Wolfenden—namely, the size, 1:0 mm., and the

proportionate lengths of abdomen and furca 7: 8 : 9—come very

near °97 mm. and 21:23:28, the corresponding measurements

in my specimen. The lateral prolongations of the third and

fourth thoracic segments are short in both specimens. In

Wolfenden’s specimen, however, the spine on the second joint of the

second antenna is said to be ‘“‘not more than half the length of

that of the Ist basal, and shorter than the distal hook,” but in

mine the spine in question is just one-third as long as that on

the Ist basal and is distinctly longer than the distal hook.

Cleve (4) has described a species, C. erythraus Q@, from the

Red Sea, which is very closely allied to both the above-mentioned

forms, but if his description be taken as accurate it must be

distinct. The female genital segment has sete on the genital

openings and a ventral hook, the endopodite of the 4th foot has

two setze, and the furca is moderately long. ‘The size 88-94 mm.

is. slightly less than that of C. dubius. The proportions of the

— oe

FROM CHRISTMAS ISLAND. 293

abdominal segments and furea are 10:10:12, 7. e. the genital

and anal segments are equal; in the figure, however, the genital

segment is shown as being the longer. The most noticeable

difference is in the 2nd antenna, in which the spine of the 2nd

basal is figured as being half as long as that of the 1st basal, and

is said to reach to the distal margin of the joint. In my species

the spine of the 2nd basal is only one-third as long as that of the

Ist, and falls short of the distal margin of the joint by half its

own. length.

In view of this uncertainty it seems advisable to give the

Christmas Island specimen a distinct name -until the specific

characters of C. amazonicus and C. erythreus ave more fully

known.

Female (Pl. XIV. fig. 5)—Length 97 mm.; cephalothorax in

mid-dorsal line ‘58 mm.; abdomen °39 mm. (EIS XY, fie. 6) ;

proportions of abdominal segments and furca, dorsally 21 : 23 : 28,

ventrally 16:22:27. Sides of cephalon parallel, Ceph. + Th. 1

being about twice as long as broad. Points of 3rd thoracic seg-

ment (Pl. XIV. fig. 7) not very long (broken in my specimen),

with thickened margin on posterior edge. 4th thoracic segment

short, with sharp points. Genital segment about 14 times as long

as wide, with sete on genital openings, in lateral view with a

small sharp hook anteriorly on ventral edge. Anal segment a

little longer than genital segment and slightly overlapped by it

dorsally, about three times as long as wide. Furea slightly

diverging, a little longer than anal segment. Furcal rami about

1x10, terminal setz missing.

1st antenna six-jointed, proportional lengths of joints : - ae

2nd antenna (Pl. XIV, fig. 8) with spine on Ist basal very long;

spine on 2nd basal short, about 3 as long as that on 1st basal and

falling short of the nearest point of distal margin of joint by

about half its own length. Distal margin of 2nd basal with two

sharp teeth. Terminal spine of 2nd antenna very strong, shorter

than spine on 2nd basal.

Mouth-parts not satisfactorily made out.

Jointing and number of sete of swimming fect normal.

Ist foot : exopodite with broad terminal spine, almost as long as

endopodite, with fine denticulations.

2nd foot (Pl. XIV. fig. 9): terminal spine of exopodite almost

straight, with moderately coarse denticulations on outer edge and

two small distal serrations on inner edge (possibly an individual

eharacter and not specific).

3rd foot with long slender terminal spine on exopodite, the

proportional lengths of joints and spine being 28 : 20: 60: 70.

4th foot (Pl. XII. fig. 7): endopodite small, with two setee ;

- exopodite slender, the proportional lengths of its joints being

approximately 6:4:5; outer edge seta on Ist joint almost

as long as 2nd joint, blant process on 2nd basal joint internal to

endopodite.

Proc. Zoou. Soc.—1911, No. XX. 20

294 MR. G. P. FARRAN ON COPEPODA

CorycHUS ANDREWSI, sp. n. (Pl. XIII. figs. 7-9, Pl. XIV.

figs. 1-4.)

Female (Pl. XIV. fig. 1).—Length -95 mm.; cephalothorax in

middle line ‘64 mm., to end of thoracic spines “7 mm.; abdomen

-31 mm. (Pl. XIV. fig. 4). Proportional lengths of abdominal

segments and furca 19:12:10. Furcal rami about five times as

long as wide. The genital segment is about 3 as wide as long

and 4 as deep, in lateral view, as long; the antero-ventral angle

is very marked in lateral view, being almost a right angle,

rounded off at the apex and furnished with a small patch of

minute spinules. The points of the 2nd thoracic segment are

short and notched on the inner margin. The points of the 4th

thoracic segment are very short, but sharp at the extreme tip.

The eyes are of medtum sizé and are separated by rather less

than their own diameter.

1st antenna, proportional lengths of joints ae:

2nd antenna (Pl. XIII. fig. 9): Ist basal with long spine; 2nd

basal with spine about $ as long as that on the first, and two

rather slender teeth on its dista! edge. The terminal spine of the

2nd antenna is about half as long as that on the 2nd basal.

The proportional lengths of the jomts of the swimming-feet are

best seen in the figures.

The outer edge spines of the exopodites of the 1st and 2nd feet

(Pl. XIII. fig. 8, Pl. XIV. fig. 3) have well-developed laminz, but

on the 38rd foot they are very slightly laminate. The terminal

spine on the exopodite of the 2nd foot is slightly eurved, on the

ist and 3rd feet it is straight. In the 4th foot (Pl. XIII. fig. 7)

the outer edge spine of the Ist joint of the exopodite is short and

slender. The endopodite of the 4th foot bears two moderately

large sete.

Five specimens, females, were found in two gatherings.

Comparing OC. andrewsi with the other species in which there

are two sete on the endopodite of the 4th foot, we find that it is

separated from C. lautws, C. lwhbocki, C. tenuis, C. erythreus,

C. venustus, C. anglicus, and C. africanus by its short furea,

which isless than 4 of the length of the rest of the abdomen.

In OC. amazonicus the anal segment is said to be longer below

than the genital, a fact which at once distinguishes that species.

In C. asiaticus and C. minimus, according to Dahl's table, the

furca agrees in being about half as long as the genital segment,

but the genital segment is said to be respectively two and three

times as long as the anal, while in C. andrewsi it is only 14 times

as long. @

CorycUS MURRAYI, sp.n. (Pl. XIIT. figs. 1-6.)

Female (Pl. XIII. figs. 1, 2).—Length 1-2 mm.; cephalothorax

in middle line :75 mm.; abdomen and furca -43 mm. The pro-

portional lengths of the abdominal segments and furca in‘01 mm.

are 18: 13: 12. The wings of the 3rd thoracic segment

FROM CHRISTMAS ISLAND. 295

(Pl. XIII. fig. 3) are long, sharp-pointed, and spreading, the

width between the points being % of the width of the cephalon

and equal to the length of the abdomen without furea. The

abdomen is wide in front and regularly tapered to the furea,

the segmentation between the genital and anal segments being

rather obscure. Immediately behind and exterior to each genital

opening is a small blunt process or papilla projecting beyond

the margin of the genital segment in dorsal view. ‘The set

at the genital openings are very small and might easily be

overlooked.

The 2nd antenna (PI. XIII. fig. 4) is much stouter than in

C. andrewsi and has one strong tooth on the inner margin of the

2nd joint. The seta on the 2nd joint is rather more than half as

long as that on the 1st joint, and reaches almost to the end of

the inner edge tooth. The terminal claw of the 2nd antenna is

strong.

The swimming-feet have no special characters. The terminal

spine of the exopodite of the 2nd foot (Pl. XITT. fig. 6) is only very

slightly curved. The endopodite of the 4th foot (Pl. XIII. fig. 5)

bears two moderately strong setz, and the outer edge spine of the

lst joint of the exopodite is rather short.

This species is rather closely allied to the above-described

C. andrewsi, having, like it, setee on the female genital segment,

two setze on the endopodite of the fourth foot, and a short furca.

Six specimens were found in three gatherings.

LITERATURE REFERRED TO.

(1) Brapy, G. S.—‘ Challenger’ Reports, vol. viii. pt. xxiii.

Copepoda, 1883.

(2) Cuaus, C.—Die freilebenden Copepoden, 1863.

(3) Curve, P. T.—“ Plankton from the Indian Ocean and

Malay Archipelago.” Konig]. Sv. Vet.-Akad. Handl.,

Bd. 35, No. 5, 1901.

(4) Creve, P. T.—‘‘ Plankton collected by Mr. Thorild Wulff

during a Voyage to and from Bombay.” Arkiv for

Zoologi, Bd. 1., 1903.

(5) Dany, F.—Verh. Deutsch. Zool. Ges. 1894.

(6) Dana, J. D.—Proc. Acad. Nat. Sci. Philadelphia, ii,

1845.

(7) Dana, J. D.—Ann. Nat. Hist. vol. xviii., 1846.

(8) Dana, J. D—Proc. Amer. Acad. Sci. ii., 1849.

(9) Dana, J. D.—U.S. Explor. Exped., vol. xiii. Crustacea,

1852.

(10) Giesprecut, W.—Fauna u. Flora des Golfes von Neapel,

vol. xix. Copepoda, 1893.

(11) Woxrenpen, R. N.—Fauna and Geography of the Maldive

and Laccadive Archipelagoes, vol. ii., suppl. 1, Copepoda,

1905.

20*

296

ON COPEPODA FROM CHRISTMAS ISLAND.

EXPLANATION OF THE PLATES.

All the figures have been drawn with the aid of a camera lucida.

Puate X.

Fig. 1. Corycella brevis 9, dorsal view.

2. 5 » @, lateral view.

3. Ms ep ordeLoot

4. i See oplstaoote

5. , » ¢, 4th foot.

6. a ©, 3rd thoracie segment, mounted.

Wo Corye ella curta °, 3rd foot, exopodite.

8. 33 9 Sy Pinal foot.

9. Bs » §, 8rd thoracic segment, mounted.

10. a & © , 2nd antenna.

11. i >» +, 4th foot.

Prater XI.

Vig. 1. Coryeella curta 9, dorsal view.

2. 5 » §, lateral view.

3. ne » 9%, abdomen, dorsal view

4. * x a, abdomen, dorsal view.

5. Bh » 6, Ath foot.

6. » 6, 8rd thoracic segment, mounted.

@. Cor cella brevis 9, 2nd antenna.

8. Corycella sp., 6, dorsal view.

8), a 3 6, 4th foot.

10. Corycella carinata 2 , abdomen, lateral view.

ll. Coryceus gracilicaudatus 6, dorsal view.

12), & 55 6d, lateral view.

Poate XII.

Wig. 1. Coryeeus catus 9, dorsal view.

2. 3 >» ¢, srd and 4th thoracic segments, mounted.

3h Ny > £2, 4th foot.

4. Coryceus obtusus 2, 4th foot.

5. i oe 9, 3rd and 4th thoracic segments, mounted.

6. ©, 2nd antenna.

fo Oly yes dubius OF ‘4th foot.

8. Coryceus tenuis 9, 3rd and 4th thoracic segments, mounted

9. 5 » @, 2nd antenna.

Prats XIII.

Fig. 1. Coryceus murrayi 2 , dorsal view.

2. 7 5 ©, lateral view.

3. os ey ©, 3rd and 4th thoracic segments, mounted.

4. ss 5 9, 2nd antenna.

B, s 5 9, 4th foot.

6. 5 * 9, 2nd foot.

7. Coryceus andrewsi &, 4th foot.

8. 5 be ®, 2nd foot.

9. i $3 2, 2nd antenna.

Prats XIV.

Vig.1. Coryceus andrewsi & , dorsal view.

2. ks Ss ©, 3rd and 4th thoracic segments, mounted.

3. ss x 2, 1st foot.

4. ©, abdomen, dorsal view.

5. Corycceus dubius 2, dorsal view.

6. 5 >» &, abdomen, lateral view.

ie 5 >» @, ord and 4th thoracic segments, mounted.

a » @, 2nd antenna.

» §, 2nd foot, exepodite.

ON SOME NEW ZEALAND SPIDERS. 297

14. On some New Zealand Spiders.

By H.R. Hoaa, M.A., F.Z.8.

| Received November 11,1910: Read February 7, 1911. |

(Text-figures 92-96.)

Prof. Chas. Chilton, of Christchurch, New Zealand, suggested

to his students that they should in the vacation collect specimens

at various points over the North and South Islands, wherever

their homes were situated. The result is that from some half-

dozen localities specimens of spiders were brought in within the

space of a few weeks, and these he has kindly sent to me. ;

New Zealand for a considerable period has been rather well

worked from a zoological point of view, and its spider fauna

described not only by Dr. Koch but by local men such as

Messrs. Urquhart and Goyen. It is therefore not a little re-

markable to find in the small collection here described so large

a proportion of new species, and, moreover, the same new species,

in one case from three, and another from two widely separated

localities at the same time. The thirty specimens comprise no

less than twelve species of eleven genera. Of the species, four

are new, and one a local variety of an Australian species not

hitherto recorded from New Zealand. They were collected mostly

from Ruakura and Wellington, in the North Island, by Miss B.

D. Cross, and one by Mr. E. P. Turner from the summit of

Mt. Ngauruhoe shortly after an eruption. The latter appears

to have minute particles of volcanic dust adhering to it.

The species from the South Island were taken by Messrs.

R. N. Hawkes and G. M. Thomson, and by Prof. Chilton himself.

On Mt. Peel, in the Province of Canterbury, Mr. Hawkes

picked up at random a male Uliodon of a new species, a new

species of dArgoctenus, a male of Cambridgea. antipodiana (with

its unique form of stridulatory organ), and three females of Por-

rhothele antipodiana Walck., a new locality for the two latter

species.

In 1905 (Zool. Jahrb. xxi. pt. 4,1905) M. Simon formed a genus

Mynoglenes for a spider (JZ. insolens) from the Chatham Islands,

500 miles east of New Zealand, and I was recently able to describe

another species (Reports Phil. Inst. of Canterbury, N. Z., 1909,

vol. i. p. 165) from the Sub-Antarctic Auckland and Campbell

Islands, south of New Zealand, but the genus had not so far been

recorded from New Zealand itself. Miss Cross, Prof. Chilton, anc

Mr. Turner send specimens, one each from separate localities—

Ruakura and Mt. Ngauruhoe in the North, and Picton m the

South Island. These, although differing in size, are all adult and

appear to be similar. I have taken them to be the same species.

Hemiclea rogenhofert L. Koch and Araneus verrucosus Walck.,

from Ruakura, are among the few spiders hitherto known to

be common to both New Zealand and Australia ; and Tetragnatha

298 MR. H. R. HOGG ON SOME

v4)

ferox Li. Koch, with some slight variations, is now for the first

time recorded from New Zealand.

Prof. Chilton sends Desis marina Camb., taken by himself in

Lyttelton Harbour, and a Pardosa from Picton Harbour.

Suborder MYGALOMORPH 2.

Family AVICULARIDA.

Subfamily DIPLURINA.

Group MACROTHELEZ.

Genus PoRRHOTHELE Simon.

PoORRHOTHELE ANTIPODIANA Walck,

Three females from Mt. Peel, Canterbury. Collected by

Mr. R. N, Hawkes,

Suborder ARANEA VERA. Section ECRIBELLATE.

Family DRASSIDAL.

Subfamily HEMICL@IN4,

Genus Hemicitaa Thor,

HEMICL@A ROGENHOFERI L, Koch,

One male from Ruakura. Miss B, D. Cross,

Family ARGIOPIDAS.

Subfamily ARGIOPIN&,

Group 7'eTRAGNATHE#,

Genus Terracnatua Latr,

TETRAGNATHA FEROX L. Koch, var, CROsS#, nov.

One male and two females from Ruakura. Miss B. D. Cross.

A few variations from the Australian form are noted below,

but it is so like in other respects that I do not think it can be

specifically separated therefrom,

Group ARANEE,

Genus Meta Clerck,

Mera arcenrata Urquhart.

Nephila argentatum Urq. Trans. N.Z. Inst. vol. xix. p. 92.

One male and two females from Ruakura. Miss B. D. Cross.

These correspond so nearly in all respects with Mr. Urquhart’s

description of the Vephilaabovementioned, except that the side eyes

NEW ZHALAND SPIDERS. 299

are close together, that I have no doubt as to their belonging to

the same species. The silvery colouring of the abdomen is

suggestive of some of the Vephile, especially V. venosa L. Koch,

but the shape of its cephalothorax and shorter lengths of tarsal

joints show that it is not a Nephila, of which genus no species

have been recorded from New Zealand.

Genus ArAneEus Linn.

ARANEUS VERRUCOSUS Walck.

Two males and six females from Ruakura. Miss B. D. Cross.

Family CLUBIONID&

Subfamily CTENIN&.

Group CTENEA.

Genus Uxropon L. Koch.

ULIODON HAWKESI, sp. n.

One male and one female from Mt. Peel, Canterbury.

Mr. R. N. Hawkes.

One male from Wellington, Miss B. D. Cross.

Subfamily LIocRANIN &.

Group Mi7TvrRGE«.

Genus Arcoctenus L. Koch.

ARGOCTENUS AUREUS, Sp. Nn.

One female from Mt. Peel. Mr. R. N. Hawkes.

Family AGALENIDA.

Subfamily Cy B H1N &.

Group Desizz.

Genus Drsis Walck.

Desis MARINA Cambr.

One female from Lyttelton Harbour. Prof. Chilton.

Group CYBZE.

Genus MyNoGLENEs Simon.

MYNOGLENES CHILTONI, sp. n.

One male from Ruakura. Miss B. D. Cross.

One male from Mt. Ngauruhoe, Waikato, N. Island. Mr. EK. P.

Turner. Shortly after an eruption.

One female from Picton, 8. Island. Prof. Chilton,

300 MR. H. R. HOGG ON SOME

Group ARGYRONETE#.

Genus CAMBRIDGEA.

CAMBRIDGEA ANTIPODIANA White.

One femalefrom Mt. Peel, Mr, R. N. Hawkes.

Family PISAURIDAL.

Group DOLOMEDE,

Genus Dotomepes Latr,

DOLOMEDES TRIDENTATUS, Sp, n,

One female from Dunedin, Mr, G. M. Thomson.

Family LYCOSIDAH,

Group PARDOSE.

Genus Parposa C, Koch,

(Probably) P,. canescens Goyen,

Lycosa canescens Goyen, Trans, N.4Z, Inst, vol, xix. p. 203.

The specimens, one male and one female, agree closely with

Mr, Goyen’s description of his species, which he states he always

found on river banks. Prof, Chilton collected this from a mud-

bank in Picton Harbour, and I have little doubt as to its being

the same, Its short broad lip, square front, two large equal

teeth and one smaller on the inner margin of the falx-sheath, and

procurved front row of eyes, show clearly that it belongs to

C. Koch’s genus Pardosa, and this would follow from Mr. Goyen’s

description of the lip alone,

TErRAGNATHA FEROX L, Koch, var. cRoss#, nov, (Text-fig. 92.)

Female, Cephalothorax, mandibles, maxille, cox, and legs

orange-yellow. The lip, except on the front edge, is yellow-brown

and the sternum somewhat darker. The scanty hairing is rather

pale brown, and the spines on the legs dark brown, The cephalic

part of the cephalothorax is bordered with a somewhat browner

line, but with white hairs thereon, and there are two pairs of

yather darker spots on the front of the rear slope. The abdomen

above is yellow-grey, bordered with a dark grey wavy line on each

side, the whole distance from front to rear, There are short white

hairs rather thicker on the side slopes than on the back. Under-

neath it is a dingy yellow-grey with short brown hairs; there is a

yather darker area running from the spinnerets to the breathing-

apertures and epigyne.

The cephalothorax is a long oval, not quite twice as long as

broad, rounded at the sides, the cephalic part distinctly raised above

the thoracic. Between the end of the former and the rear slope

NEW ZEALAND SPIDERS. 301

Text-fig. 92.

Tetragnatha ferox, var. cross@. Male.

_ a, eyes; 6, mandible of male from inside.

C, . female from inside.

d, rf male from outside.

e, lip and maxille; f, epigyne; g, male palp.

302 MR. H. R. HOGG ON SOME

are two pairs of depressions, one behind the other, divided by a

rather broad ridge.

Viewed from alnove both rows of eyes are recurved, ihoee of the

rear row equal and equidistant two and a half times their diameter

apart. The front median eyes are nearer together than the rear,

and apparently rather larger; they are situated on the sides of a

protuberance, and are as far from the rear eyes as from one

another. The respective pairs of side eyes are a similar distance

apart. As they are all dark and on black rims, the real size of

the eye is difficult to distinguish. The clypeus is as broad as a

front median eye. :

The mandibles are as long as the cephalothorax, divergent, and

protrude at an angle of 45 degrees. They have long upstanding

hairs on the inner side and shorter ones on the outer edge. On the

outer margin of the falx-sheath there are five rather long teeth,

the upper one on the top edge being the largest. On the inner

margin there is one large tooth at the top, one at the corner, then

one small one a little way down, and then seven nearer together

reaching to the top of the lip. On the inner side of the fang itself,

near the base, is a short stout spine midway between the two

marginal spines on the top edge of the sheath.

The mawille are long, broadest at the anterior end, where they

rather bend back with a rounded corner at the outer edge; from

here they slope in to about the middle and then out again. The

trochanter of the palp is very long.

The lip is about two-fifths the height of the maxille, rounded

in front with a rather prominent edge. It broadens out towards

the posterior end, which is also rounded, and there isa depression

all the way round a little distance from the margin.

The sternwm is a long narrow shield-shape, twice as long as

broad, hollowed in front, and with a prominent apophysis at each

of the front corners ; it is smooth and shiny, with a few fine up-

standing hairs. There is a space between the second and third

pair of coxe, and the fourth pair are not quite contiguous to

one another.

The abdomen is long and narrow, straight at the sides and

slightly rounded at the tr ont and rear.

The epigyne is a transverse oval, the inside filled with an oval

convexity.

The palpi are thin. The tibial joint is twice as tone as the

patellar.

The legs are long and thin ; the tibia of the front pair more than

six times as long as the patella, There are four pairs of spines at

the sides of tibie 3 and 4; the spines are fine and not very long.

The male is coloured the same as the female, except that the

sternum is of the same bright yellow colour as the cephalothorax,

maxille, and legs.

The mandibles are quite as long as the cephalothorax; on the

inner margin of the falx-sheath are twelve teeth, the second of

these being the largest. On the outer side there are three large

NEW ZEALAND SPIDERS. 303

protuberances at the end, and five rather large teeth at intervals

down the margin of the falx-sheath.

The male differs from 7’. ferow L. Koch in the larger second

tooth from top on the inside of the mandible, in the apophyses at

the head, the two inner curving in the opposite direction to the

outer, in the mandibles being quite as long as the cephalothorax,

instead of shorter, in the tibial joint of the palp being distinctly

longer than the patellar, and the palpal stigma being not more

than half as long as in 7’. feroxw, and the legs longer in proportion.

The measurements (in millimetres) are as follows :—

Male.

Long. Broad.

Cephalothorax ... 3 Dy

Migomen 6. ...0 5s 1

Mandibles ...... 3

Trochanter Patella Metatarsus

Coxa. & femur. & tibia. & tarsus.

Me Ssy acy a ee 9 10 ee 3 ll

2 Il if 7 7 = 22

oe 1 3s 24 3 = 10

A 1 63 64 6a 204

Baliov sis, ane = 24 2 lS = 64

Female.

Long. Broad.

Cephalothorax ... 3 12

Abdomen.......... u 2

Mandibles......... 3

Pat. & Metat.

Coxa. Tr. &fem. tib. & tars.

egsy.cersat i vaeei G 8 iat & tr45

Deeks 6 6 6 = 19

3 il 23 2 24 = 8

4 1 5 5 a 16s

Balin nesenec eee = 24 2 — 4

T. ferox L. Koch, fairly common along the eastern and southern

coasts of Australia, has not been previously recorded from New

Zealand. The New Zealand and Australian females are practi-

cally identical, and I consider this to be a local variety, which i

have named after Miss Cross, who sends a male and a female from

Ruakura.

ULIODON HAWKESI, sp. n. (Text-fig. 93.)

Female. The cephalothorax is bright, rather pale yellow-brown,

with short upstanding brown bristles and downlying white hairs.

On the cephalic part there are three longitudinal darker lines—one

median reaching from the eyes to the rear end, and one shorter

each side, also a darker line round its margin. At the sides of the

thoracic part is a darker brown marginal stripe.

304 MR. H. R. HOGG ON SOME

Mandibles dark yellow-brown, with downlying white hairs and

upstanding brown bristles; fangs bright red. Lip and maxille

yellow-brown, with greyish-yellow hairs. Sternum yellow-brown,

with whitish hairs. Coxe pale yellow, with a dark band at the end.

Femora yellow, with three dark bands underneath. The other

joints of the legs dark yellow-brown, with black spines; grey

scopule on all tarsi and metatarsi.

Abdomen above greyish-yellow mottled with black, sparsely

covered with brown upstanding bristles and downlying white hairs.

On the under side the fore part is yellow, with short yellow-brown

hairs as far as the pulmonary apertures, the remainder pale yel-

lowish grey. The sides are darker, nearly black, with longitudinal

lines of yellow and white hairs.

Text-fig. 93.

th /

—

Sth hi a ye ,

See SS

Uliodon hawkesi. Female.

a, eyes; b, male palp; ec, lip and maxille; d, epigyne.

The cephalothorax is convex, highest at the beginning of the

rear slope, thence downwards to the eye-space. The cephalic part

is rather more than half the greatest width of the thoracic, its sides

rather straight, but those of the thoracic rounded. A long deep

longitudinal fovea reaches to the top of the rear slope, which is

steep.

The rear row of eyes viewed from above is recurved, but

procurved from in front; the median eyes their diameter apart

and rather more than three times that distance from the laterals,

which are of the same diameter. Of the shorter front row, the

laterals are as large as those of the rear row, but the median are

NEW ZEALAND SPIDERS. 305

two-thirds their diameter. These are all equidistant, the diameter

of the median apart, and one and a half of the same from the

corresponding eyes of the rear,row.

The elypeus is the width of the front median eyes to the margin

of the behaired part, and there is an equal distance of smooth

muscle to the insertion of the mandibles.

The mandibles are much kneed at the base, narrowing to the

anterior end, thickly covered with bristly hairs. The fangs are long

and powerful. On the inner margin of the falx-sheath are three

large equal teeth, and one Jarge between two smaller on the outer.

The dip is longer than broad, hollow in front, rounded at the

sides, and constricted at the basal part, which is rather long. It is

more than half the length of the maxille, which are narrow atthe

base and gradually wider to the upper margin, where they are

broadest, curving gradually on both inner and outer sides.

The sternwm is a broad shield-shape, slightly convex, pointed at

the rear between the nearly touching coxe. It is thickly covered

with upstanding bristly hairs.

The legs are stout and moderately long. The trochanters have

a median ring of bristles rising from a chitinous ridge. The

tibie and metatarsi have long powerful spines, with a bunch

at the anterior end of the latter. There are thick scopule on the

tarsus and metatarsus of all legs. The two claws have five or six

pectinations. '

The palp of the female has a pair of long spines at the base of

the distal joint ; the claw is smooth.

The abdomen is ovate, pointed at rear end, and the spinnerets

terminal. The inferior pair are conical, two-jointed, the second

joint bemg short and hemispherical. The superior pair are

cylindrical, half the length of the inferior, with similar second

joints.

The measurements (in millimetres) are as follows :—

Female.

Long. Broad.

Cephalothorax ... 8 Z fi EEO

Abdomen ......... U1 7A

Mandibles......... 33

Pat. & Metat.

Coxa. Tr. &fem. tib. & tars.

IDES Rapa 1 2 63 7 t+ = 99

2 2 54 63 bs = ae

3 2 6 De 6 = 192

Ye 63 7 Sees Von

TEI CT lah eae etna if 34 2 2 9

One male and one female collected by Mr. R. N. Hawkes on

Mt. Peel, Canterbury, at an altitude of 2000 feet.

One male from Wellington, collected by Miss B. D. Cross.

306 MR. H. R. HOGG ON SOME

Besides being twice as large, this species differs from Uliodon

(Zora) frenatum L. Koch, from the same province, in the longi-

tudinal lines on the cephalothorax. It has no pattern on the

abdomen, and the sternum is yellowish instead of black-brown.

There is a scopula on the tarsus and metatarsus of all legs,

instead of only on the front two pairs.

ARGOCTENUS AUREUS, sp.n. (Text-fig. 94.)

female. The cephalothorax is bright yellow in the middle

the whole distance from the eyes to the rear; this is bordered on

the side slope with dark grey, almost black, reticulations, with

parallel stripes of the same colour reaching to the margin. At

Text-fig. 94.

Argoctenus aureus. Female. X 2.

a, eyes; 6, epigyne; c, lip and maxille.

the margin itself, on the rear half, is a narrow band of yellow,

and at the side of the eye-space is rather darker yellow. The

whole is covered with short downlying white flat hairs, the rear

fovea being black-haired. Behind the eye-space is a jet-black

curved line passing on each side along the margin, and there are

NEW ZEALAND SPIDERS. 307

two black patches on the elypeus, but the same thick white

hairs. The mandibles are deeper yellow, with a black streak on

the inner edge of each, reaching from the base almost to the point,

where it is again yellow. ‘The fangs are red-brown. The lip is

ervey, the maxillz pale yellow, with broad nearly white margins

on the inner side of the former and front of the latter. ‘The

sternum is dark yellow-brown, thickly covered with white hairs

and grey bristles. The legs are pale yellow on the upper side,

with a grey streak nearly the whole length of the under side on

the femoral, patellar, and tibial Jomts ; covered with white or pale

greyish-yellow hairs. The scopule and claw-tufts are pale grey.

‘he abdomen is yellow-grey, with a brown median scolloped

stripe, darkened at the edges, reaching from the base to the

spinnerets. The whole rear part is rather darker with a reticu-

lation of grey on the yellow, the stripe remaining darker still.

The hairs are white, with the exception of a few upstanding

brown ones on the stripe and at the base. The under side is

greyish yellow, covered with white intermixed with short brown

hairs. The hairs on the yellow spinnerets are brown.

The cephalothorax is convex, rounded at the sides and broadest

a little in front of the rearend. The front part is about one-half

the greatest width, and the sides of this, as far back as the rear

row of eyes, are straight.

‘The eyes are in four rows. The second row, the laterals of the

front row, being vertically just clear of the median first pair.

These are all of the same diameter and half their width apart.

The rear two pairs are again of equal size, one-third larger than

the front, the median ones being one-third of their diameter apart

and the same distance from the front laterals. The median are

14 times their diameter from the rear laterals respectively, and

the latter are four times their diameter apart.

The clypeus is as broad as the front eyes.

The mandibles are kneed at the base, divergent, and at the

anterior end half their width at the base.s They areas long as the

front of the cephalothorax is wide. There are two teeth on the

inner margin of the falx-sheath, and three smaller on the outer.

The sternum is broad shield-shaped, straight in front and

curving toa point between the rear coxe, which are not quite con-

tiguous. It is thickly covered with downlying flat hairs and

upstanding bristles.

The lip is broader than long, rounded at the side and in front,

where it has a broad sloping edge.

The maille are convex, upright, and about three times the

height of the ip. From their base they curve outwards on both

inner and outer margins to a rounded upper end, so that they

are broadest in the middle. They have a few short upright

bristly hairs.

The legs are moderately fine, covered with long powerful spines.

There is a thick scopula on the tarsus and metatarsus, and claw-

tufts under the weak curved pair of claws, On tibie 1 and 2

308 MR. H. R. HOGG ON SOME

axe three pairs of spines on the under side, on metatarsus of same

a pair of very long spines at the base underneath, and another

about the middle at the side. On tibia 4 are three long spines on

the upper side and three pairs on the under. On metatarsus 4

are two pairs on the under side, three long spines on the outer

side, and three long single ones on the upper side.

The tibia and patella of the palp are of equal length.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax ... 34 | in Ero nt

Alodomienti ease e. D4 4

Mianidiileseeeerese: ls

Pat. & Metat.

Coxa. Tr.&fem. tib. & tars.

Megs. 1 Le 4 A 45 = 14}

2 les 4 4} oe Sy id

3 1 3 33 + ie wlll

4 14 4) 5 Oo)

Baie ce tecs aacre: 2 2 2 WR 6

One female from Mt. Peel, Canterbury, collected by Mr. R. N.

Hawkes.

This species would appear to be somewhat near M. Simon’s

A. vittatus from Noumea, but the median abdominal stripe

extends the whole length, and is pale brown instead of intense

black. The legs are more heavily bespined, and the femora,

although streaked longitudinally, are not ringed with black.

MYNOGLENES CHILTONI, sp. n. (Text-fig. 95.)

Male. The cephalothorax is of a rather deep yellow-brown,

darker brown in the striations, but paler again on the rear slope.

The eyes are pale yellow. The mandibles bright yellow-brown

with red fangs. Lip and maxille about the same as the man-

dibles. The sternum darker, and dingier, is covered with fine

granulations. The legs are pale yellow all over, with fine brown

hairs paler towards the extremities.

The abdomen, both on the upper and lower sides, is black-brown,

scantily covered with fine recumbent brown hairs.

The cephalic part of the cephalothorax is slightly raised above

the thoracic and rounded in front, where it is one-half the width

of the broadest part, the latter being two-thirds of its total length.

The surface of the thoracic part.is smooth, without hairs, very

finely granulated. The central fovea is shallow and round, with

four pairs of small lateral furrows, and a broader and deeper pair

round the margin of the cephalic part, all radiating from the

fovea.

The eyes of the rear row are equal, the median their diameter

apart, and the laterals one and a half diameters therefrom ; these,

with the front laterals touching them, lie on prominent tubercles.

NEW ZEALAND SPIDERS, 309

The front median eyes, half the diameter of the others, are one-

half of their own diameter apart; they are as far from the rear

median as the latter from the side. The clypeus is as broad as

the median eye-area is long, and the deep indentation characteristic

of this genus runs below the front row of eyes.

The mandibles are much swollen in the upper half, whence

they narrow and become almost cylindrical in their lower half ;

they are as long as the front of the head part is broad. On the

outer margin of the falx-sheath are two long teeth followed by

two smaller.

Text-fig. 95.

Mynoglenes chiltoni.

a, female; b, male; ec, eyes; d, male palp from the front; e, male palp from

the side; f, epigyne; g, lip and maxillze.

The lip is broader than long, round at the top, with the pro-

truding upper margin of the genus, and less than half the height

of the maxille. These are upright, small at the base and gradually

widen out as they curve over the lip.

The sternwm is a broad shield-shape, straight in front, rounded

at the sides and running to a point at the rear, where the cox

do not quite meet. The latter are rather unusually large and

rounded, especially the anterior pair.

Proc. Zoot. Soc.—1911, No. X XI. 21

310 MR. H. R. HOGG ON SOME

The trochantal joints of the legs are long, and have a pronounced

protruding ridge round them, about the middle. The other joints

are thin and tapering, the tarsus and metatarsus being quite

fine; they are well covered with very fine greyish-brown hairs,

but have no spines.

The abdomen is long and oval, the surface being much corru-

gated longitudinally. The spinnerets are short, with a small

hemispherical second joint.

Except in size the female agrees with the male.

The measurements (in millimetres) are as follows :—

Larger Male.

Long. Broad.

1 in front.

Cephalothorax ... 3 | 5 in front

INicdlomenwe seers 4. 2,

Mandibles ...... ils

Pat. & Metat.

Coxa. Tr. & fem. tib. & tars.

WWegsis asncce ters 3 3 a ee OL

: : 3 a

3 Zz 22 2 a = +

4 2 8) i A. = 102

Pa aie ctounceiy es 4 ii 2 3 = 35

Female.

Long. Broad.

1 in front.

Cephalothorax ... 2 { ; ie er

Abdomen ......... 2 12

Mandibles ...... 1

Pat. & Metat.

Coxa. Tr. & fem. tib. & tars.

ue gs yet ue - 2 23 Ht 2 = . i

SW gee be Th Nh ELE

Je 2 2 _—

Ae is 12 2 = 5

Pall snece sees z 1 1 - = 3

This species differs from M/. marrineri in the paler colouring of

the mandibles, lip, maxille, and legs. Thereis no yellow pattern

on the back, only a slight mottling, and no paler area on the under-

side of the abdomen as in the foregoing. The thoracic fovea is

short and shallow instead of being long and deep, and the

granulated portions are confined to the thorax and sternum.

One male from the summit of Mt. Ngauruhoe, Waikato, N.

Island. Mr. H. Phillips Turner.

One male from Ruakura. Miss B. D. Cross.

One female from Picton in the South Island, found by Prof.

Chilton, after whom I have named the species.

NEW ZEALAND SPIDERS. SL

DoLoMEDES TRIDENTATUS, sp.n. (Text-fig. 96.)

Female. The cephalothorax is a bright chestnut, a broad area

behind the eyes being thickly covered with silvery grey hairs,

from this a narrower median streak of similar hair reaches to the

rear slope, on each side of this is a broad area of chestnut

covered with short orange hairs, and thence nearly to the side

Text-fig. 96.

Dolomedes tridentatus. Kemale.

a, eyes; 6, profile; c, mandible from inside; d, tarsal claws; e, epigyne.

margin is again a broad white-haired streak. The mandibles are

dark red-brown, thickly covered with long bristly hairs, nearly

white on the outer sides, darker grey on the inner. The fangs

are red. The lip, maxille, and sternum are dark brown, thickly

covered with long upstanding brown hairs. The legs and palpi

are bright orange, with long brown hairs darkening towards

21*

Sle MR. H. R. HOGG ON SOME

the extremities of the limbs. The muscle parts are bright white.

The abdomen is bright yellow-brown, thickly covered with short

dingy yellow-brown hairs, a paler median stripe reaching from the

base to about halfway down the back ; on the sides of the abdomen

the hairs are white. The underside is quite dark yellow-brown,

with four longitudinal white-haired rather narrow stripes. The

spinnerets are bright yellow-brown.

The upper surface of the cephalothorav although flat is con-

siderably raised above the level of the side margins, and slopes

steeply at the rear. It is one-fourth longer than its breadth im

the widest part, and the front is only half the width of the

latter. It is rounded at the sides and rear. There is a long and

deep median longitudinal fovea extending from the cephalic part

to the rear slope.

The side eyes of the front row are on the lower slope of black

tubercles, which, extending farther back than the median eyes, give

the appearance from above of a recurved row. Their lower edges

are on a level with the lower edges of the median, and as they are

smaller the line of the upper margins is really procurved. The

median eyes are about their diameter apart and one-half that

distance from the side pair, which are two-thirds of the width of

the former. The eyes of the second row are about twice the —

diameter of the median eyes of the front row. They are rather

less than their diameter apart, the same distance from the front

median, and the total length is clearly less than that of the front

row. ‘The median eye-area is broader than long. The eyes of

the third row are four-fifths the diameter of those of the second,

five of their diameters apart, and two of the same from the eyes

of the second row. The clypeus is about 25 times as broad as the

front median eyes.

The mandibles are kneed at the base, stout, as long as the

front of the cephalothorax is broad, and slightly divergent. ‘They

are thickly covered with long upstandmg bristles. The fangs

are broad at the base, strong, and well curved. About the

middle of the inner margin of the falx-sheath are three large

teeth, all equidistant, at the lower end of the outer margin

is one large tooth between two small. The lp is broader than

long, straight in front, and less than half the length of the

maxille.

The sternwm is a broad shield-shape, straight in front, pointed

at rear, where it passes between the two rear coxe, which are not

quite contiguous.

The legs are stout and straight, slightly and normally tapering

to the anterior ends, but in nowise fine or flexible; they are

armed with numerous long and strong spines. There are scopulee

on the tarsi and metatarsi of the front two pairs, on the tarsi

only of the rear. The superior claws have six or seven teeth, the

inferior are smooth. There is a pair of spines on patelle of pairs

3 and 4, none on patelle of 1 and 2. The femoral joint of the palp

NEW ZEALAND SPIDERS. oily

is incurved and enlarged at the anterior end, the tibial joint is

longer than the patellar, and the female palp-claw has four pecti-

nations.

The abdomen is oval, rounded at the sides. The hairs are

smooth and fine. On the upper side are three pairs of muscle-

spots. The epigyne is horseshoe-shaped, hollow in the anterior

half only, and without any longitudinal rib. The superior pair

of spinnerets is longer than the inferior. They are both two-

jointed, the second joint half the length of the first.

The egg-sac is globular, white, and flocculent, 12 millimetres

in diameter.

The measurements (in millimetres) are as follows : —

Long. Broad.

Cephalothorax ... 8 | ; oat

Abdomen ......... 13 64

Mandibles......... 3

Pat. & Metat.

Coxa. Tr. & fem. tib & tars.

Theos sae il 3 7 a 63 = 234

Bae 7 7 64 = 234

eee od GE Gu =. Ot

Aen s3 7 8 8 = 2%

1ERT1/ Se eee me oe 3 ee etm

One female from Dunedin. Collected by Mr. G. M. Thomson.

This species differs from the accepted limits of the genus

Dolomedes as detailed by M. Simon in having three teeth only

on the inner margin of the falx-sheath instead of four. The

clypeus while broad is less broad than the area of the median eyes,

and the patellee of legs 1 and 2 are without spines. It is, however,

much nearer to Dolomedes Latreille than to Thaumasia Perty,

Drances Simon, or any other genus. I would have felt it right

to propose a new genus for it on the above differences but for the

fact that it graduates so imperceptibly into other members having

four teeth, notably D. hattoni and D. trippi from the Chatham

Islands, which are certainly Dolomedes. Its eyes and clypeus, the

equality of the Ist and 2nd pairs of legs, and the marking of

the cephalothorax in the female correspond almost exactly with

those of D. trippi. Its epigyne is of the same type as that of

D. huttona.

314 MR. A. H. COCKS ON A

EXHIBITIONS AND NOTICKS.

February 21, 1911.

Dr. A. Smrra Woopwarp, F.R.S., Vice-President,

in the Chair.

Tur Secrerary read the following report on the additions made

to the Society’s Menagerie during the month of January last :—

The registered additions to the Society’s Menagerie during the

month of January were 121 in number. Of these 63 were

acquired a presentation, 17 by purchase, 32 were received on

deposit, 5 in exchange, and 4 were born in the Gardens.

The total number of departures during the month, by death

and removals, was 160.

Amongst the additions special attention may be called to the

following :—

2 Lion Cubs (Felis leo), from the Sebakwe River, 8. Rhodesia,

presented to the King’s South-African Collection, through

H.R.H. The Duke of Connaught, K.G., by Col. Weston Jarvis.

Deposited by H.M. Tur Kine on Jan. 14th.

1 Jaguar (felis onca), born in the Gardens on Jan 24th.

2 Bay Duikers (Cephalophus dorsalis), from Coomassie, pre-

sented by Capt. 8. H. Chapin on Jan 21st.

2 Virginian Deer (Dorcelaphus americanus) 2 2, from North

America, purchased on Jan 17th.

1 Nacunda Nightjar (Podager nacunda), captured at sea off

the coast of Brazil, new to the Collection, purchased on Jan 2nd.

~1 Bornean Fireback Pheasant (Lophura ignita), presented by

H.G. The Duke of Bedford, K.G., Pres.Z.8., on Jan 21st.

‘Dr. H. Hammonp Suir, M.R.C.S., F.Z.8., exhibited three

skins of male Pheasants assuming female plumage, sent to him

by Mr. Arthur Gilbey, and some microscopical specimens of the

glands prepared by Dr. 8. G. Shattock.

Mr. G. A. Boutencer, F.R.S., V.P.Z.8., contributed a paper

based on a collection of Fishes from the Lake Ngami Basin,

Bechuanaland, made by Mr. R. B. Woosnam, F.Z.8.

This paper has been published in the ‘ Transactions.’

Mr. Aurrep H. Cocks, M.A., F.Z.8., exhibited a series of

photographs of the female Brindled Gnu recently born in the

Society’s Gardens, and gave the following account of its growth

and coloration :—

“On receipt of the orphan calf of the Brindled Gnu, born in

}

‘

YOUNG BRINDLED GNU. 315

the Gardens on the 1st December last, which was brought to me

on the 3rd, I was struck by the great difference between her and

the drawing by J. Smit in P.Z.8. 1900, Plate xlviii. (facing

p- 771), representing another calf previously born here; that

figure, and one in ‘Illustrations of the Zoology of South Africa,’

&e., by Andrew Smith, M.D., London, 1849, No. 16, Aug. 1842,

Plate xxxvii.*, being, I believe, the only two existing of a Gnu

calf of this species.

Young Brindled Gnu born in the Society’s Gardens on December 1, 1910.

From a photograph taken on January 7, 1911.

“What seemed specially worthy of note was that the tail for

the proximal two-thirds of its length was white, as if the white-

* The beginning of A. Smith’s description of the young is quaint: “ Form and

appearance clumsy and unseemly ”!

316 MR. A. H. COCKS ON A

tailed species were the older form. The body-colour was also

quite different from that shown in Smit’s plate, being of the lustre-

less ash-brown of an immature house-mouse, with a dull or rusty

black dorsal stripe terminating in a point at the sacrum. The

cheeks instead of being smooth and matching the body in colour,

as shown in both the above-mentioned plates, were exactly the

reverse. The dark colour on the outer side of the ears did not

Text-fig. 98.

Head of young Brindled Gnu.

From a photograph taken on January 7, 1911.

quite reach the margins, as if a man had taken a brushful of

paint, and had given one streak to each ear, without afterwards

making good the deficiencies. The lower two-thirds of all four.

legs was quite white; Andrew Smith’s plate shows this con-

spicuously. The whole face was very dark or black, the black

extending to a sharply defined width of about half an inch round

the lower side of the eyes.

YOUNG BRINDLED GNU. Ply

“ By the 13th December (that is, when the calf was twelve days

old), the whole of the upper side of the body was a light fawn (the

original ‘puppy coat’ having been shed); the legs were gradually

colouring, or less pure, or conspicuously white, than at first; and

the colour was extending from above downwards. The white on

the tail was already nearly gone. Face dark; nose and muzzle,

as before, black; cheeks quite light, the dark ring remaining

round the eyes. The coloration of the exterior of the ears as

before.

“Tn Smit’s plate the fawn of the body is darker, and more

mixed with black-tipped hairs, and the drawing was evidently

made at a greater age* than the present example attained to,

not only for the reason just stated, but because the tail-tuft is of

a brighter black, the dark circle round the eyes is shown as the

faintest possible shadow, and the general figure of the animal

is more set; but the most important discrepancy—which seems

only explicable on the supposition that the present example be-

longed toa local race widely differing from the two calves previously

figured,—is that both plates show quite a smooth cheek, whereas

the present example, like her father (and I feel sure the mother

was the same), had a perfect and very conspicuous ‘ Newgate

fringe,’ consisting of a woolly moustache, like a poodle’s, but

situated about halfway along the gape, and quite distinct from

the moustachial bristles on the muzzle, meeting bushy whiskers

on the cheeks, and continued as a long thick beard on the chin ;

the whole cheek was hirsute, or covered with long hairs projecting

in a more or less upward direction.

“By the 16th the horns were appearing through the skin, and

she had cut some molar teeth, which we could hear her grinding,

so we tied up a wisp of hay, and she very soon learnt that it was

good to eat. The tail now much resembled that of the Grey

Squirrels so familiar in the Park, with the addition of a black tuft.

“By the 27th the dark ring below the eyes was fading

away.

“On the 14th January the horns were about an inch long ; and

on the lower surface of the chin, between lip and Newgate

fringe, but hardly extending to the side face, was a white patch,

which I had not previously noticed.

“On the 15th the calf appeared as vigorous and in as perfect

health as ever, but on the morning of the 16th she lay prostrate,

and in a very short time was dead; the cause, as decided by

Mr. Plimmer, the Society’s pathologist, being broncho-pneumonia.

“The Gnu was photographed, by myself, on December 13, on

December 31 and January 2, by Press photographers, and on

January 7, by request of Dr. Chalmers Mitchell, by a Henley

photographer ; the series to some extent shows the growth.

* No hint is given (Joc. cit.) as to the age of the calf when drawn, but it was born

en July 14, and the drawing exhibited on November 20.

318 MR. H. G. PLIMMER ON DEATHS

“T took the following measurements immediately after death :—

Extreme length, tip of nose to tip of tail (= vertebre), 4ft. 13 ins.

Length of face, 114 ins.

ear, 52 ins.

‘3 tail (to end of vertebrae), 11 ins.

i fore leg (elbow to end of toe), 227 ins.

" hind leg (hock to end of toe), 154 ins.”

PAPERS.

15. Report on the Deaths which occurred in the Zoological

Gardens during 1910. By H. G. Prrmmer, F.R.S.,

F.Z.8., Pres.R.M.S8., Pathologist to the Society.

[Received February 6, 1911: Read February 21, 1911.]

On January 1, 1910, the number of animals in the Zoological

Gardens was 3186, and during the year 2354 animals were

admitted, making a total of 5540 for the year.

The number of deaths during the year has been 1554, that is

a death-rate of 28 per cent.; but if from the above total we

deduct 643 animals which did not live for six months after their

arrival in the Gardens—that is, roughly, the time at which we

find they have either got over their journeys, or died from any

disease they brought with them, or have got entirely used to their

new enyironment—the percentage of deaths is reduced to 16:4,

which is less than that of the last two years.

The following figures will show the general relations of the last

three years :—-

1908. 1809. 1910.

Total number of animals ......... 5608 5303 5540

Totalideaths eee eet eceercerre SM 1492 1554

Percentage of deaths ............ 31 28 28

Percentage of deaths, excluding

those which occurred in ani-

mals which had noé been six

months in the Gardens ...... 17 17°8 16-4

Tf we consider the weather conditions of 1910, and compare

these figures (bearing in mind the relative number of animals),

it will be seen that the percentage of deaths of 1910 is more satis-

factory than that of the two preceding years.

The total deaths are divided as follows: Mammals 362, Birds

845, Reptiles 347.

The following tables show the facts ascertained in outline.

Table I. sets forth the actual causes of death in each of the three

groups specified. Under Reptiles are included Batrachians and

Fishes.

ile

IN THE GARDENS DURING 1910.

Tasie I.—Analysis of the Causes of Death.

Disease.

Microbic or Parasitic

Diseases.

Mulberculosisteres sesh eee se ene

My COSISIe ny ee at k= za een

[RCO MA eye ears is ees See

Septiceemia ag Sea ne ee

LAS QDS), Gad S68 282k Ae Omens aie

EAL ELUTbISHY trees A eee cs Seesa chi tate

1DVTA OA ASTED” Gao ssocoenacnoaaecneecese

iRericanditises-esseec ssa eee

ReNIOTILEISt eee yates ee eee

Stomatihisueces: cones eee

Hydatids

Worms Kor ee eet ER Ee.

Hemogregarines - ee Re ene

Malaria # aa Rap HdaeAGHeR Rn eee ae

@oecidirosiseey eee ee

Leucocytozoon infection .........

Qystibispes eee he cece:

Pancreatitis

Diseases of Raspindirey

Organs.

Bronchititsts: cot ete ee ae:

Broncho- -pnewmonia . Pe AEE

Congestion of lungs ...............

Atelectasis Behtetbere tae et etme ve

. Diseases of the Heart.

Dilatation of heart ...............

Fatty degeneration

. Diseases of Liver.

Fatty degeneration ...............

Hepatitis...

Diseases oltay Tract. |

Over-disterision of crop .........

Gastritis .

Gastric ulcer: ation .. aes:

GCOSIRO-QMTUIS 54,000 condaatoonaooe:

IBniterib Secs aoe eee

Intussusception....... 0.0.2.2...

Intestinal obstruction ............

. Diseases of the Urinary and

Reproductive Organs.

INephinibisness sees rea eat ake

Stone .......

Inflammation of oviduct .

. Diseases of Nervous System.

Myelitis

. Various.

Carcinoma .

Sarcoma ..

Puerperal eclampsia ... Rae

Senile tdecaiya sa. ess eee

PRICK Cts ce etya seas eee ee

Leukemia ........

Anemia without ascertained 2

GAMISO Ms eccaioecseereeee net aees

Injuries discovered post-

mortem steeoe ofe }

Mammals.

me co 0

Bw AONHREA

DR wor

319

Reference

S : to Notes

Birds. | Reptiles. following.

163 9 1

99 6 2

73 82 3

eae =

3 5

D) 3

“ - 7

eee 16

eae 1

1 9

eee Soe, 10

112 14 oe

15 1

4, if

wae 1

3 10 12

186 27 ie

ed wee 14

eae 15

eee 16

5 /

1 18

16 2

320 MR. H. G. PLIMMER ON DEATHS

Besides those tabulated above,

113 animals were killed by order or by companions,

101 3 died of starvation and malnutrition,

7 » Were not examined,

108 4, ~~ ~+were too stale for detailed examination,

these completing the total.

In Table I. the classification is made into those diseases which

actually caused the death of the animals, but in a large number

of Mammals, Birds, and Reptiles other lesions were present which

helped towards the fatal issue, and these are classified in Table II.

If taken together with Table I. a much more accurate estimate of

the amount of disease in the Gardens will be arrived at.

Taste II. —Other Diseases found in the Animals tabulated in

Table I.

Reference

Diseases. Mammals.| Birds. | Reptiles. fo Notes

Mulberculosis: se wccsen nee cc eee eon

NEV COSIS HES eee eeecerrace cence eR eras

Pneumonia ee eae eee ee oe

Stomvatibis ieee eee eee ree enn) aeaenn

Worms See ae rae ora eos ee

(Peritonitisie ees acsce ene Oee ee eee eens

Pericarditis es Nice hi a ache Aine

I Bh dah oS} G0 megane suaccadoesdanedimpe conse sed adosds ae =

ARTAATTNOSOGNES ooo oon conode vce ada ote oncde aac ie 2 i 19

JE EAC UOINES. sco 000 cs van cncace vce aoa csv cos ate ae 1

Baneneatitisy a Saceqecs, sec ceeiccee a eee ee As 7

IBIGTIRIbISH ERE mses ceeen ee One eee eee ae

CAYEL aT Ge He nepngareonebosncbadaunmanonesene advert ase

i 0 Gb) oe

eS bo

er)

jx)

Bronchitis ee eons eee ee

Bron Ch O-pnewmMOnlaleeeee ee ee eeeee eee eee

Congestion of lungs .............6.000...000 0

(Clea OVE MWOWETS “Goons ass ogconscas ooo coosse cus

(Edema of lungs Langue Semod aeilatiogn ate

JE GHCIROW MORES? Sooscc cde dooonsaa9 00 00006 noe 000 oar

Wika ted@hearh mscntcsecte osaseiec se ccee conchae

Fatty heart BPE Seen aRAG Ona ae

Hlatitiyg liver Snenceaveee cea erase eee

Hepatitis

Grastiitisieecccr ae aeons are oe eee

Gastric cenailoneeseet pee ena eeeteee

Gastno-entenitise ene a eee eee: a

WRbeRIbIS pers eae eG ek ee 12

Intestinal obstruction ........................ aa

INephritisiaer.: Se cot ee ct eae 13

UNS CIDES anaes caterer soren ecei-n hon nas shan cone we bee

Osteo-anthini hist net asate otc es ae ereee il bce ABs 20

AUC KIGES ore vnr ants arene PR aR Cs ceree ci ores eae 23 le

ee bo

Gre 0

He:

(oon

—

OD: MOA Oe oo bo om Ob

pes

> g

Me wow ca:

no:

TE oc:

IN THE GARDENS DURING 1910. ayAll

Taste I1I.—Shows the Distribution of Diseases causing Death

amongst the principal Orders of Mammals.

Diseases.

Mycosis

UMM benGwlOsiseneese tine ice cesiedecu vaaeee

IPT CUMON ae eee eee te aera cont eee

Se phiGre miley ayeeee-ces- che ocecccetest. sae

JPISDIBIING) A= seater Coo aee RB anEEAne EER eeerE Bot

JN OREGEDS SaBsce ABR Gab eee naneen cnet meme bees

WTS CTE caategasnenera Gas cee aL ARDC See

IRERICALGIbISe- e-eee e ee

Peritonitis

i SIsrGlE Na aa he ed Le Sots A

WSAGTET IC) p BIA a ame ec” BES A aeRO ale ARORA

Oystitishe.chnsacdndee

Primates.

Carnivora

Rodentia.

Marsupialia.

Ungulata.

Edentata.

Bronchitis

Broncho-pneumonia

Congestion of lungs

Atelectasis

Di lataiony ot heantierrcersse copes esse

Tint i\y¢ 0200 oes copeepochne rice seal ececs errs

Baby ALiVet ws ssieuast lek Woden. x2) asad

Eve paititiispcesatre eee cet cone ec cae eects Sane

Gastritis

Gastric ulceration

Gastro-enteritis

Intussusception

Myelitis

Sarcoma

Senile decay

OLAS cs ao tate aaa

INE LIEISIa Ld uae aaNet

RS LON Cree OGORRBOCE boa th Gena sonncee ae

(Sirsa V7 gi soi a aR

IBC WAT NSE): neann cboeebpaade oom ioboaaosnned sat

TRG) RUS Seoacd aaaudeenSHanasmpuepbedbcuseeteer

TENT SS i eooeadaau seeceas ca MaueAasan ase sod ee

fico: ~T

bo ove

bore:

be orm \)

(o2)

wa :

TH be:

bo: OV: Gow:

HOrwH:

os On oo

ees:

ro oo:

bobo sy:

As Tuberculosis and Mycosis have again been the cause of a

large number of deaths amongst the Birds, the following Table

has been drawn up to show the relative incidence in the various

Orders.

Taste [V.—Showing the Comparative Incidence of Tuberculosis

and Mycosis in the various Orders of Birds.

Passeres

Orders.

PRT CANT ces Seidel ee ee ee

PSIG ACTS is cAdet, Anat eae Ae

PASTIS GLOSS oc -hase dee

@ohomibser esas ee eee

(Gealliinias! Sas ee ae. ee

URID ES) sc dv ec. ase eterna aN

anid aes 42, ee ee ee ee

Sinwthronese yay eee eet

Tuberculosis.

Mvcosis.

(a)

ON DEATHS IN THE GARDENS DURING 1910.

Notes on the foregoing Tables.

The following Notes refer to some points of special Ane

which it was not possible to include in the tables :—

1. It will be noticed that there has been about the same

percentage of tubercle as last year amongst the Mammals and

Reptiles; that is, it has been found in 10 per cent. of the

Mammals and 4:9 per cent. of the Reptiles examined. In the

Birds, however, there has been an increase of 107 cases; that is,

it has been found in 24°9 per cent. of those examined. This

increase amongst the Birds has been partly due to an epidemic

which occurred in the New Bird House in the beginning of the

year. This was dealt with by entire disinfection of the house,

and some structural alterations which have greatly improved it ;

a very definite improvement has taken place, but the arrange-

ment of the cages at the ends of the house, and overcrowding

generally amon ost the Birds, is a constant source of danger.

Amongst the Mammals 2 cases have been of bovine type, and

amongst ‘the Birds there have been 7 cases of this type.

The liver of a Zantalus showing this particular type has been

sent to the Museum of the Royal College of Surgeons.

2. Mycosis (under which term several distinct mould-diseases

are grouped) has also increased among the Birds to a considerable

degree—107 cases as against 48 last year.

The Mammals dying from this disease have been principally

Wallabies and Gazelles. The disease (formerly grouped under the

Septiceemias) has been found to be caused by a mould which enters

through the mouth and produces abscesses about the jaws and in

the adjacent muscles, and eventually a septicemia.

Tt has also been found that a disease of the eye in Birds is due

to a mould. It commences as a keratitis, and then pus is

formed in the anterior chamber, and secondary abscesses form,

from which the bird dies. Of the division headed Reptiles

3 were fishes with a mould-disease of the skin which invaded the

muscles.

3. The increase of pneumonia amongst the Reptiles (87 cases as

against 64 last year) has been mainly due to the variable

temperatures in the Reptile House, owing to alterations in the

heating apparatus. Only 3 of these cases were due to the presence

of worm-eggs and embryos. There has been a decrease amongst

the mammals and birds.

4. Five of these cases followed injuries: the remaining two

were in Wild Swine, where it was caused by an organism belong-

ing to the group producing Hemorrhagic Septicemia.

5. The one Mammal was a Fox with suppuration in the middle

ear and destruction of the semicircular canals.

6. In an Antelope the pericarditis was due to numbers of

worm-cysts on the visceral pericardium, which, so far as I know,

is unique. In the Birds it was mostly due to the deposition of

uric-acid crystals in the pericardium,

ere xeti

y Nty-

‘ v6

iuler:

d.Green delet Chrom

WILOS A.

—

Vd £7

ma tyr

for

var teentala.

ON THE VARIATIONS OF THE SPOTTED SALAMANDER. 323

7. In the Mammal—an Aardwolf—the worms were encysted

pentastomata, which were in large numbers all over the body,

causing pleuritis and peritonitis. In the Birds they were mostly

syngamus.

8. Under Malaria are grouped cases in which intracorpuscular

parasites belonging either to the Halteridium or Proteosoma

group have been found in sufficient numbers to cause death.

9. In a Bulbul in which 60 per cent. of the polynuclear leuco-

cytes were infected. This I believe to be new.

10. Most of these occurred during the first and last three

months of the year ; 4 of them were badly rickety.

11. Nine of the Mammals had bad rickets also. The number

of cases amongst Birds, in which it is relatively much more fatal,

is slightly less than last year.

12. In most of the Mammals it was caused by food-poisoning,

in others, and in the Birdsand Reptiles, by worms burrowing into

the mucosa.

13. In 7 Mammals, 59 Birds, and 1 Reptile the enteritis was

hemorrhagic; in6 Birds and 6 Reptiles it was due to worms; and

in 9 Birds and 1 Mammal it was due to foreign bodies. This

disease is a little less prevalent than last year.

14. Two very extreme intussusceptions occurred in two Wom-

bats, recent arrivals, which came together.

15. Ina Coypu Rat, in which 27 stones were found.

16. Four of these cases of cancer were in Wallabies, the

stomach in all was the seat of the primary growth; the fifth was

in a Markhoor in the mouth.

17. Two Gazelles died from sarcoma, one of liver and one of

mediastinal glands.

18. This occurred in a Jungle-fowl and was of the spleno-

medullary variety.

19. In a Partridge and Marsh-Bird, both not described before.

20. In a Bat, in which all the wing-joints were affected.

16. A Contribution to the Study of the Variations of the

Spotted Salamander (Salamandra maculosa). By

Epwarp G. BouLencEr*.

[Received December 10, 1910: Read February 21, 1911.]

(Plate XV.T & Text-figures 99-102.)

The experiments now being carried out in Vienna by

Dr. Kammerer on the colour-changes of the Spotted Salamander

(Salamandra maculosa) in relation to its environment are

attracting attention, and it has occurred to me that a general

survey of what is known of the varieties of this very variable

species, especially in connection with the geographical distribution,

* Communicated by G. A. Bourznecer, F.R.S., V.P.Z.S.

+ For explanation of the Plate see p. 347.

324 MR. E. G. BOULENGER ON THE

would not be without importance at the present moment. Such

a survey would afford those who wish to follow the path opened

up by Dr. Kammerer a more precise basis than can be obtained

from the available literature on the subject. I have therefore,

with the help of my father, undertaken a revision of the rich

material in the British Museum and have now the honour of

offering an account of it for publication to the Zoological Society.

One ‘of the principal results of my study has been to lay greater

stress on the disposition of the markings than on their actual

size, form, or colour, and to define two principal forms in Central

Europe, which previous authors have not separated with sufficient

precision, in spite of their well-marked geographical segregation.

Except in the case of var. molleri, with its aberrant coloration,

authors dividing S. maculosa into a number of subordinate forms

have dwelt on real or supposed structural characters, whilst

ignoring the disposition of the markings. Bedriaga (8, p. 98),

the most recent writer on the subject, recognizes, a part from the

typical form, also called by him var. ewropea (2, p. 252), three

varieties, namely—var. algira, var. corsica, and var. molleri, the

two former based only on slight structural differences, which are,

besides, not constant, as J intend to show further on. Under the

designation of typical form authors have generally thrown together

specimens with different styles of markings, either simply ob-

serving that these are subject to infinite variation or classifying

them under a number of titles, which refer merely to indi-

vidual variations, such as the var. teniata, var. quadrivirgata and

var. nigriventris, proposed by Diurigen (10, pp. 577 & 578) for

certain individuals. In order to avoid introducing a new name,

T will adopt the first of these for the assemblage which I have

endeavoured to define and contrast with the typical spotted form

on which the name maculosa is based.

To better bring out the individual differences to which the

markings are subjected in this Salamander, I have drawn up

tables of a certain number of the specimens in the British

Museum, upon a scheme which should prove of use to those

making experiments on the colour-changes, as by that means a

record of each individual specimen, out of a large number, can be

kept in such a way as to ensure its future identification. Such

tables, explained by the annexed diagram (text-fig. 99), do not,

however, convey an exact representation of the markings, which

can only be done by descriptions, but they will be found to answer

well enough for the purpose of identification.

In defining the varieties into which the species Salamandra

maculosa may be divided, I have not lost sight of occasional

exceptions, and have duly pomted them out. There are always

exceptions, especially when we have to deal with forms of sub-

specific rank, but such as I have come across are too few to

militate against the adoption of a var. teniata as opposed to the

forma étypica.

In the tables the explanation of the various columns is as

VARIATIONS OF THE SPOTTED SALAMANDER. 325,

follows :—The length of each specimen (in millimetres) is taken

from the end of the snout to the posterior extremity of the vent.

O means presence (+) or absence (—) of the supraocular spot

(on the upper eyelid).

P, the spot on the parotoid gland.

Text-fig. 99.

Diagram to explain the tables of

different markings in Salamandra

maculosa.

S, spot or spots on the snout.

OP, OS, whether the above-

mentioned spots are confluent or

not. PB, whether or not the

spot on P is confluent with one

on the body. In cases where the

two sides differ, the initials R

(right) and L (left) indicate the

discrepancy.

Sp., total number of spots on

the upper surface of body (to

a line connecting the posterior

borders of the hind limbs), which

is divided into four conventional

segments, numbered I, II, III,

IV; under each of the latter

figures, the spots pertaining to

each segment are given. These

spots are numbered 1, 2, 3, &c., in

the order shown on the diagram ;

in case of absolute symmetry, the

spots are numbered from right to

left.

A stands for the vertebral area,

bearing the two median rows of

glands. B (right and left) for the

area between A and C; C (right

and left) for the area occupied by

the series of large lateral glands ;

D for the area between the latter

and the limit of the belly (taken

from an imaginary line connect-

ing the lower border of the axil

with the groin). In column D,

right and left sides are separated

by a —, the figures referring to

the number of spots on each side

(spots confluent with the ventrals

and dorsals not reckoned), not to

the individual spots as in the

columns A to C.

G, presence (+) or absence (—) of yellow spots on the chin and

gular region.

V, on the ventral region of the body.

Proc. Zoou. Soc.—1911, No. XXII.

MR. E. G. BOULENGER ON THE

The tabulation of the markings on the specimen represented in

text-fig. 99 may be drawn up as follows :—

SECTION. AREA.

foe SS pe ES ce a Se

O OP P PB S OS Sp. I. I. Ml. iV. C B A B mG

+ — + — — — 6 1128 45 6 —12.4.6 1.2.4.6 1.3.4.5.6 3

The following is a hst of the specimens preserved in the

British Museum and the Lataste Collection, arranged geographi-

cally under varieties :—

ForMA TYPICA.

France.

ils Gis Beure, near Besangon. Mme. Phisalix.

Oks hs Avignon. M. M. Mourgue.

By Basses-Alpes. M. Honnorat. Lataste

Collection.

4. gf. S.E. Franee. Prof. Duboseq.

Corsica.

5-9. 9 & yg. Bocognano. Dr. J. de Bedriaga.

10. Yg. Vizzavona (1100 m.). Prof. Vayssiére.

ll. Ye. Corsica. Dr. J.de Bedriaga. Lataste

Collection.

Germany.

12. ¢. Near Stuttgart. Prof. K. Lampert.

4 (see. Balienstadt, H. Harz. Dr. W. Wolterstorff.

14-15. S$. Sharfenberg, near Meissen, us

Saxony.

Switzerland.

16-17. ¢ 2 Near Lausanne. W. Morton, Esq.

18. Her. Fliielen, Uri (465 m.). M. A. Ghidini.

19. g. Lugano, Ticino (275 m.). a

20. g. Val Bavano at Cavergno, ee

Ticino (ca. 600 m.).

Ttaly.

21. ¢. Prego, Brianza, L. Como. Dr. C. Vandont.

We, 2. Tuduno Olona, Varese. Be

23-24. 9. Olginate, Prov. Lecco, s

Lombardy.

25-54. ¢. Cascinella, near Borgoli, Prov. Dr. R. Gestro.

Genoa.

55. 3. Viterbo. Prof. J. J. Bianconi.

56-60. ¢ 2. Prov. Rome. Prof. Carruccio.

61. 2. Aspromonte, near Reggio, Prof. O. Neumann.

Calabria (1600 m.).

Austria-Hungary.

62-63. ¢ 2. Hiitteldorf, near Vienna. Dr. F. Werner.

64-66. ¢ 9 & yg Near Prague. G. A. Boulenger, Esq.

67. 2. Brasso, Hungary. Prot. L. v. Méhely.

68. Ye. Teszla, Bozau Mts., Hungary. oF

69-88. gQ. Nagy Beeskerek, Hungary. Hr. A. v. Kovacs.

89. ¢. Hungary. ;

90. 2. Sarajen, Bosnia. Dr. F. Werner.

91-92. g. Travnik, Bosnia. 56

VARIATIONS OF THE SPOTTED SALAMANDER.

Roumania.

UBL Gee

94-113. $ Q& yg.

Near Azu,

Moldavia.

Sinaia, Carpathians of Vallachia.

Carpathians of

Greece.

114. Yg. Parnassus.

Asia Minor.

ley, & Zebel Bulgar Dagh, Cilician

Taurus (1200 m.).

Algeria.

yg. Mt. Edough, near Bona,

119-122. ¢2 & yg. A ip

116-118. 2 &

123. [oe 9 2

124. 9. L’Arba, near Algiers.

WS ée Algeria.

Morocco.

126-133. 92 Benider Hills, near Tangier.

Vars. GALLAICA and MOLLERI.

Spawn.

lp Qs Vigo, Galicia.

2-5. 62 “yg. Galicia.

6-11. g2 & yg. Loroya Valley, near Madrid

(800-400 m.).

12. Yg Spain.

Portugal.

18) 2, Coimbra.

14-17. g8 rY)

18-19. df Cintra.

WO, YX Near Lisbon.

Var. TANIATA.

France.

Near St. Malo.

We We

2-5. dQ. Roscoff, Finistére.

6-8. 2 & yg Near Rouen.

OQ Qe

LON OF Armainvilliers, near Paris.

11-13. ¢. Haute-Marne.

14-19. g 9. Beure, near Besang¢on.

20. 2. Bourg-en-Gironde.

21-26. ¢ 2. Hérault.

ile, Nise Aix-les-Thermes, Ariége.

28.3). Eaux-bonnes, Basses-Pyrénées.

Belgium.

29-30. f Maredsous, Prov. Namur.

ees Waulsort, i

M. A. Montandon.

Dr. T. Kriiper.

C. G. Danford, Esq.

Dr. Hagenmiiller.

“Lataste Collection. .

M. L. Bedel. =

M. Lallemant. ,,

M. H. Vaucher.

M. V. L. Seoane.

“Lataste Collection.

M. de la Escalera.

Lord Lilford.

Dr. J. de Bedriaga.

Col. Yerbury.

Sr. Mattozo Santos.

G. A. Boulenger, Esq.

M. Louis Miller.

M. L. Horst.

M. E. Simon. Lataste

Collection.

Dr. A. Pettit.

Mme. Phisalix.

Lataste Collection.

Prot. Duboseq.

M. V. Baillet.

Rey. F, A. Walker.

Rey. B. Lebbe.

G. A. Boulenger, Esq.

29%

328 MR. E, G. BOULENGER ON THE

Luxemburg.

Sy, @- Luxemburg. M. V. Ferrant.

Germany.

53-40. 62 & yg Tisenberg, Harz. Dr. W. Wolterstorff.

41. 2. 5 * W. H. Decks.

42-51. 62. Harz. Zoological Society.

62-53. Ye. 53 G. A. Boulenger, Esq.

64-55. 6. Vorwohle, Brunswick. Dr. W. Wolterstorft.

56-58. ¢ 9. Holzminden, Brunswick.

59-60. 6 @. Stadtoldendorf, Brunswick. M. A. Ghidini.

61-63. ¢ & yg. Lippe-Detmold. Dr. J. Roux.

64-67. 62 & yg Near Stuttgart. Prof. K. Lampert.

68. ¢. G. Duchy of Baden. Basle Museum.

Switzerland.

69-72. 62 & yg. Langenbruck, Jura. Basle Museum.

73. 3. St. Gallen. M. A. Ghidini.

Portugal.

74-75. 6 & yg. Oporto. KE. Allen, Esq.

76-77. 6. Portugal. P. B. Webb, Esq.

I. Tue TypicaL Form.

This form deserves to be regarded as the typical, not only

because the name maculosa applies best to it, but because it is the

only one found in Austria (ef. Werner, 32, p. 119), where it was

deseribed under that name by Laurenii (18, pp. 42,151). It also

happens to be the form figured by most authors :—Aldrovandi (1,

p. 641), Résel (24, frontispiece), Latreille (17, pl. i.), Sturm (30),

Reider & Hahn (23), Funk (12, pl. 1.), Bonaparte (5), Rusconi

(25, pl. i.), and Camerano (6, pl.i.). It corresponds to the vars.

A and C of Dumeéril and Bibron (9, p. 37), a—c of Schreiber (27,

p. 75). Inthis form the black nearly always greatly predominates

over the yellow, the latter appearing as markings of various

shapes,—round, elongate, C- S-, Y-shaped, d&c., and disposed over

the body, often in 3 to 5 alternating series, or with a median

series forming a sinuous or zigzag vertebral stripe. If, as 1s very

exceptionally the case, the dorsal spots appear to form two longi-

tudinal series, it will be observed that they by no means hang

together in regular chains continuous with the spots on the

parotoids. Only in one specimen (from Lausanne) have I felt

embarrassed as to the form to which it should be referred. Upper

eyelid and parotoid usually entirely, sometimes only partially,

yellow, the spots on the parotoid may even be entirely absent

(specimens from Algeria and Morocco). The two spots thus

located are, as a rule, distinct, but may occasionally run together.

In a specimen from Nagy Begskerek, Hungary, the yellow

markings on the upper eyelids extend across the interorbital

region, forming a cross-bar. The sides usually bear spots, which

may number up to 15, in which case they are, of course, very small.

A spot above the angle of the mouth is absent in only about 5 per

—

VARIATIONS OF TILE SPOTTED SALAMANDER, 329

cent. of the specimens. -Gular region with a few large or small

markings, sometimes completely inspotted. Below entirely black

or with : spots which are usually of small size, the only specimens

with very large spots being single ones from Genoa, Viterbo, and

Reggio in Italy. Snout generally entirely black. Limbs black,

with a characteristic yellow blotch on the upper surface of the

arm and thigh, near the base, which is constantly present; a

second on the forearm and leg may or may not also be present.

Hand and foot entirely black or with one or two (rarely more)

yellow spots, which, as a rule, are not confluent with the spot on

the forearm and leg. Single or paired spots on the tail, which

may run together to form a stripe. Under surface of tail nearly

always black.

Text-fig. 100.

Forma éypica.

The colour of the bright markings varies from chrome-yellow to -

a deep orange. I have found traces of claret-red on the head in

some specimens from Genoa. The usual absence of yellow on

the snout has been mentioned above ; the only exceptions I have

noted are to be found in specimens from Lausanne, Avignon,

330 VARIATIONS OF THE SPOLTED SALAMANDER.

Corsica, Genoa (7 specimens out of 40), the province of Rome, and

Calabria. The latter is remarkable for the large size of the yellow

markings on the upper and lower surfaces, where they almost equal

in extent the black area. The opposite extreme occurs in a spe-

cimen from the Val Bavano, Ticino, in which the yellow colour is

reduced to a few very small blotches on the parotoids and at the

base of the limbs. These two extremes are figured side by side in

text-fig. 100,

On careful examination of the specimens of var. algira Bedriaga

and var. corsica Savi, I have come to the conclusion that they are

not essentially different from the forma typica.

The var. algira is described by Bedriaga (8, p. 111) as having

the tail and digits longer and more slender than in the typical

form; but the specimens from Mt, Edough (in the Lataste

Collection), upon which Bedriaga based his observations, are in a

rather emaciated condition, having, no doubt, been kept in

captivity for some considerable time, and this is evidently partly

the cause of their slenderness. Although the digits and tails of

the Salamanders from Algeria and Morocco are, as a rule, longer

than in the typical form, the longest digits in the latter may be

actually longer in proportion to the length of the body. Thus,

in a specimen from Lake Como, the length of the longest toe is

94 per cent, of the total length (from tip of snout to posterior

end of vent), while in a specimen from Mt. Edough, Algeria, the

length of the same is 9 per cent,, and in one from the Benider

hills, Morocco, as low as 64 per cent, Again, the length of the

tail in var. algira ranges from 65 to 81 per cent. of the length of

the body, against 54 to 78 in the forma typica, an overlap which

precludes the character being used as diagnostic,

In the same author’s description of the Corsican variety, the

head is stated to be remarkably broad, and the toes to be much

more strongly depressed and with sharper edges on the sides than

in the typical form, I have examined the very specimen described

by Bedriaga, but do not find the head to be any broader than in

some of the typical and striated forms, and although the toes are

more depressed than is generally the case, they are not more so

than in certain specimens from Vienna, Bosnia, Luxemburg, and

the Harz Mountains. As to the more sharply edged sides of the

toes, this sharpness simply coincides with the degree of depression.

The supposed difference in the palatine dentition, on which

S. corsica was founded by Savi (26), has long ago been disposed of

by Schreiber (27), Bedriaga (2), and Camerano (6).

The habitat of the typical form seems to be bounded to the

west by the Hrz Mountains, the Danube, the Alps, and the

Rhone, all the specimens from east and south of that line be-

longing to it, with a few exceptions mentioned below. All over

France, west and north of the Rhone, the var. teniata, described

further on, alone occurs (with rare exceptions from the Doubs),

whence it extends to Northern Spain (Bilbao, fide Bedriaga) and

Portugal (Oporto, Brit. Mus.).

——

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mor,

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Text-fig. 101.

All the specimens hitherto examined from Belgium, North-

Western Germany, and the Rhine are referable to that variety,

which also prevails in Wiirtemberg (cf. Leydig, 19). Exceptions

to the above geographical division occur to my knowledge near, or

THE SporTED SALAMANDER. Forma typica.

—

VARIATIONS OF THE SPOTTED SALAMANDER. 333

not far from, the line of demarcation, as on Mt. Saleéve in Savoy,

according to M. Ghidini, but possibly also in Italy near Rome,

where, according to Duméril, the striped variety has been found

(yellow on the back, with three black stripes, and a few scattered

black spots on the limbs and belly)*. Further, a specimen

referable to the typical form from Ballenstedt, Anhalt, received

from Dr. Wolterstorff, one from the neighbourhood of Stuttgart,

received from Prof. Lampert, and six exceptional specimens

sketched by Mme. Phisalix: five from near Besangon, black

above with irregular yellow blotches on the back, and one from

Toulouse (Paris Museum), yellow above and below with irregular

black markings on the back, a very aberrant specimen. One of

these specimens, from Beure, near Besangon, has been presented

by Mme. Phisalix to the British Museum, and is listed under

the head of forma typica, whilst six others from the same locality

appear under var. teniata, thus showing that in the French

Jura the latter variety is not so completely fixed as it appears

to be in the North of France and Germany and Belgium.

Five specimens are represented on text-fig. 101 :—

a, from Varese, Lombardy, is remarkable fer its small amount

of yellow, and for its long and thin yellow vertebral stripe,

extending from the nape to over halfway down the back.

Sides and lower surface with only a few spots.

6, from the Benider hills, Morocco, with a small number of

large roundish spots on the back, has the left parotoid

entirely black.

c, from Hitteldorf, near Vienna, hasa remarkably large number

of spots, Flanks and sides of belly spotted.

d, from the neighbourhood of Prague, is a type with very

irregular markings. Flanks and sides of belly with only

a few large spots.

e, from Zebel, Bulgar Dagh, Asia Minor, has some of the spots

ring-like, the yellow markings having a round black spot

in the centre. Sides profusely spotted. Lower surface

with many spots of small size.

The specimen figured on Plate XV. is a male from Lugano,

Ticino, sent alive by M. A. Ghidini.

Il. THe VARIETIES OF THE SPANISH PENINSULA.

(Vars. gallaica and molleri.)

We have mentioned above that the var. teniata occurs in

Spain and Portugal. Specimens of that form, with the yellow

colour predominating over the black, are stated to occur in the

Peninsula by Schreiber (27, p. 78), and Bedriaga (3, p. 108)

* T am indebted to Mme. Phisalix for a sketch of this specimen preserved in the

Paris Museum. It is not unlikely, however, that the locality under which it

has been registered is erroneous, as neither de Betta (4), Camerano (6), nor Count

Peracca (in litt.) have come across the striped variety in any part of Italy.

334 MR. E. G. BOULENGER ON THE

records such a one from Bilbao. But in addition we find highly

remarkable specimens ranging from the form named var. gallaica

by Seoane (29), which is hardly separable from the typical form,

especially its North African representatives, to that named var.

mollert by Bedriaga, which approaches very closely some speci-

mens from near Genoa, from Austria, described by Kammerer

(cf. p. 342), and from Oran, Algeria, described by Doumergue™*.

My father has already proposed to unite the var. gallaica with the

var. molleri, a view in which Bedriaga (3, p. 109) could not concur,

on the ground that Seoane’s diagnosis does not at all agree with his

own. Itis, nevertheless, a fact that an almost uninterrupted series

can be traced between the two varieties, and I think it advisable,

provisionally at least, to regard them as extremes of one and the

same form, which is completely linked with the typical form.

The following is a translation of Seoane’s definition of the

Spanish specimens (var. gallaicw) :—“ Differs fromthe typical form,

among other characters, in the intense black of the ground-colour

and the small number of yellow spots, distributed over the

body.”

The three specimens from Galicia (Seowne) in the Lataste

Collection are remarkable for the very irregular, broken-up

disposition of the spots on the back, a few of which are partly

brownish and may have been edged with red or pink, in a manner

similar to Bedriaga’s var. molleri. The snout, supraocular and

interorbital regions, partly yellow, partly reddish brown ; yellow

on the throat somewhat predominating over the black ; sides

irregularly spotted with yellow. The number of spots on the

dorsal region of these specimens is 9, 13, and 26 respectively. A

specimen from Cabanas, Galicia (Seoane), preserved in the Paris

Museum and of which a sketch has kindly been made for me by

Mme. Phisalix, approaches very closely the Portuguese var.

mollert. ‘The British Museum specimen from Vigo differs, how-

ever, from all the above by being very scantily marked with

* Hssai sur la Faune Erpétologique de l’Oranie (Oran, 1901), p. 372.

“Corps présentant en dessus plusieurs taches jaunes et rouges sans symétrie

dont voici la distribution. Régions sus-oculaires jaunes en dessus et d’un rouge sang

en avant et en arriére. Arcades sourciliéres @un noir rougeatre. Parotides jaunes

en dessus et aussi en dessous postérieurement, entourées de noir en avant; exté-

rieurement elles sont bordées depuis l’wil jusque sur le cou, d’une longue et large

tache rouge. Seules les taches des régions sus-oculaires et celles des parotides pré-

sentent quelque symétrie. Sur le cou se trouve une grande tache transversale

échancrée en avant, 4 laquelle font suite, sur le dos, quatre taches irréguliéres (de 7

mill. sur 3 en moyenne). Ces taches alternent entre elles et touchent la double ligne

dorsale de tubercules ; elles sont 4 peu prés 4 égale distance Yune de Vautre. Prés

de V’aisselle, sur le bras, il y a une petite tache jaune bordée de rouge; une ou deux

trés petites, jaunes et rouges, se voient sur l’avant bras, et une seule sur Jes mains et

les pieds. Le fond noir des flancs est parsemé de quelques points rouges. Membres

postérieures tachés comme les antérieurs. En arriére de la ligne des cuisses, en

dessus, commence une tache jaune, longue et étroite, qui s s’étend en arriére; elle a

10 mill. sur 2 & 3. Sur Ja queue ou voit cing séries de faches doubles, rondes, qui se

rapprochent lune de lV’autre sur Ja ligne médiane supérieure; elles sont jaunes et

visiblement bien bordées de. rouge, surtout celle placées vers le bout de la queue.

Mamelon du cloaque taché de jaune de chaque cété. Dessous du corps dun violet

noiratre. Pourtour inférieur de Ja bouche bordé de taches rouges qui s’étendent sur

la gorge.

VARIATIONS OF THE SPOTTED SALAMANDER, 335

yellow, except on the gular region and on the sides of the belly.

The back, snout, and interorbital region bear no distinct markings,

but are speckled over with small yellowish dots. The parotoids

and upper eyelids are almost entirely of a reddish-brown colour.

This specimen must be regarded as an individual aberration of the

form above described, an aberration tending to the total suppres-

sion of the bright markings.

Three specimens referable to the var. gallaica have been received

from M., de la Escalera, who obtained them in the Loroya Valley,

near Madrid, at an altitude of 300 to 400 metres. The spots are

moderately large, few or moderately numerous (6 to 12), those on

the parotoids being either confluent with or distinct from those

on the upper eyelids and the dorsal region. The spot on the

eyelid in one of these specimens is entirely of a reddish brown,

that on the parotoid partly reddish, partly yellow. In this and

another specimen the interorbital region is also reddish. The

spot at the angle of the mouth in all three is brownish red in

colour. Lower surface and sides black, minutely speckled over

with yellow; throat spotted with red.

The true var. mollert is represented in the British Museum

Collection by 7 specimens from Portugal (Coimbra and Cintra),

including one of the types received from Dr. de Bedriaga, who

has thus described its coloration :—

“The colour and pattern of this variety are rather variable. The

ground-colour is usually a greyish brown, sometimes more of a

dirty grey, sometimes more brownish black or even black, broken

up above and below by pale yellow spots with an addition of grey

or greyish-brown spots into which the yellow passes gradually.

The yellow spots on the side of the body, on the hmbs, on the tail,

on the parotoids, on the throat, and on the ey alids are as if

powdered over with red dust, or washed with red, or even blood-

red. The throat may sometimes acquire a deep red colour; the

dorsal spots show here and there a red dot. The yellow spots

are very variable both in number and size; they may be either

few, in which case they are large and roundish, or numerous and

horseshoe- or ring-shaped, and forming six or eight more or less

regular longitudinal series; some of these spots break up or run

together, thus forming wavy bands. These spots may be so

numerous as to greatly reduce the ground-colour; the yellow

spots on the head are in that case the more conspicuous ‘and a

symmetrical or very ornamental pattern results.”

My father has drawn up the following notes on living specimens

exhibited in our Zoological Gardens, a few years ago.

“Some specimens were black, variegated with various tints of

grey, brown, pale yellow, and crimson. The latter colour was

particularly conspicuous on the upper eyelids, the parotoid glands,

the base of the limbs, and on the throat, but it appeared also

as small patches within the area of the more or less irregular

pale yellow spots with grey centres, which were disposed very

irregularly on the body and tail. One of the specimens was pale

336 MR. E. G. BOULENGER ON THE

olive-grey above and on the sides, freckled with black and with

pale greenish-yellow spots; the black appeared as an irregular

vertebral stripe, a dorso-lateral stripe, and bars on the flanks ;

the upper eyelids, the parotoids, and the throat were claret-red.

The coloration of such a Salamander has a lichen-like aspect

more suggestive of assimilation to the surroundings than of

warning to enemies.” Other specimens which he has seen since

had but little or no red on them, but the yellow spots were

greyish in the centre.

In his description of var. molleri, Bedriaga states that it differs

from the typical form in the snout being more pointed and

projecting beyond the lower jaw, also that the tail is shorter and

thicker. In five out of eight specimens (including Bedriaga’s

type) examined by me, I found the snout to be more projecting

than is usual in the other forms, although the most pointed

snout I have seen is in a specimen from near Meissen, Saxony.

I also found that the tail in four out of the eight specimens

was stouter and shorter than usual in the typical form and

the var. teniata, and this is also to be noticed in the figure

on Pl. XV. The length of the tail in each of the specimens

(measured from the posterior end of the vent) was 55, 55, 57,

57, 63, 67, 67, 71, the length of the body being taken as 100 (tip

of snout to posterior end of vent). The length of the tail in the

typical form varies from 60 to 78, with an average of about 67,

whilst in the var. teniata it may fall as low as 54 (Besangon).

It will be seen therefore, that there are many exceptions, and

too much importance should not be attached to this character.

I may add that the measurements of the Galician specimens,

alluded to above under the name of var. gallaica, give 57, 66,

70, and 70 as the length of the tail, and those of the specimens

from near Madrid 56, 65, and 70.

Thanks to the courtesy of the Hon. Walter Rothschild, I am

able to give a figure (Pl. XV.) of the var. mollert from a sketch

made for him by Mr. J. Green, from a female specimen in his

possession exhibited a few years ago in the Zoological Gardens.

I have myself examined two fresh specimens of this variety

from Lisbon, one alive, received from Sr. F. Mattozo Santos,

Director of the Museum Bocage at Lisbon. In the live specimen

the crimson-red colour was distributed over the parotoids, the

upper eyelids, the throat, the spots at the angle of the mouth and

on the sides, and the spot on the forearmand thigh. On the back

and tail there was no red colour, but many of the yellow spots

were partly bordered or as if washed over in the centre with a

dirty grey. In the second specimen the red colour was restricted

to the parotoids, the upper eyelids, and the spot at the angle of the

mouth. The markings of the back, limbs, tail, sides, throat, and

belly being mostly light grey in the centre and on the borders.

On careful examination of the red and grey markings in these

specimens, I have come to the conclusion that these are due, not

to special pigments in addition to or in combination with the

VARIATIONS OF THE SPOTTED SALAMANDER. Sa

yellow, as has been supposed, but to the absence of pigment, the

pigmentless flesh, highly flushed with blood, being exposed on

certain patches and the grey colour resulting i: om the absence of

yellow combined with a small quantity of black pigment.

The largest specimen of the var. mo/leri examined by me

measures 109 mm. to the posterior extremity of the vent, the

tail measuring 73.

TIL. VAR. LEANIATA.

This variety differs from the typical form in the arrangement

of the dorsal spots, which are regularly disposed in two parallel

series continuous with the patches on the parotoids, and not un-

frequently form two uninterrupted stripes. Even when the two

stripes are broken up into as many as 12 spots, these still retain

their duplex disposition, not encroaching over the black vertebral

line (Area A), or if they do so, as is rarely the case, only on the

nape and the posterior end of the body, where they may be con-

nected in H-like fashion by a cross-bar. Although in this variety

the black often predominates over the yellow, it is not uncommon

to find specimens in which the reverse takes place. In cases

where the yellow has so far invaded the upper surface as to actually

constitute the ground-colour, the black vertebral stripe may be

reduced to a mere series of spots, and Werner (81, p. 155, pl. vii.

fig. 23) even mentions and figures one in which the black is com-

pletely absent from the back. The upper eyelid and the parotoid

gland are entirely yellow (the yellow patch on the latter rarely

broken up), the two spots nearly always running together and

often also confluent with the markings on the back, which is rarely

the case with the typical form. The sides are usually devoid of

markings, and if present they only occur in small numbers.

Yellow markings are usually present on the snout, which is but ex-

ceptionally the case with the typical form. The spot situated at the

angle of the mouth is absent in about 35 per cent. of the specimens.

Gular region and lower surface of body with a varying amount of

yellow, sometimes entirely yellow with a black spot or bar on the

gular fold; the markings have often a tendency to dispose them-

selves in longitudinal series, thus the belly may be black with

a yellow lateral stripe or yellow with one or two black stripes in

the middle. Limbs with the black usually predominating ; a

characteristic yellow blotch near the base of the arm and thigh,

and a second on the forearm and leg being constant, and usually

larger than in the typical form. Hand and foot mostly black and

yellow, the yellow patches being nearly always confluent with those

on the forearm and leg. Nearly entirely yellow specimens have

likewise yellow limbs with merely 2 or 3 black spots or cross-bars.

As on the limbs, the yellow may predominate over the black on the

tail and frequently extend to the lower surface, which is rarely

the case in the typical form. The two yellow dorsal bands often

fuse on the upper surface of the tail.

The colour varies from sulphur- or lemon-yellowto a deep orange.

338 MR. E. G. BOULENGER ON THE

Some specimens with the markings almost vermilion-red, instead

of yellow or orange, obtained by Fr. v. Schweizerbath near

Stuttgart, are regarded by her as a distinct variety and named

var. coccinea (28), but this is clearly to be regarded as a merely

individual peculiarity, not deserving of a varietal name, and the

figure given by her corresponds, but for the colours of the markings,

with the var. teniata, the prevalent form round Stuttgart. Con-

sidering that the bright markings may vary, in the same locality,

from chrome-yellow to a deep orange, the so-called var. coccinea

represents merely an intensification of a tendency existing in

German specimens. Fr. v. Schweizerbath has been informed by

Prof. EK. Haeckel that such a Salamander was found by him many

years ago in the Saal Valley, near Ziegenriick, and it is not un-

likely that vermilion-spotted specimens will be discovered in other

parts of the habitat of the var. teniata. My father was informed

by an intelligent peasant woman in Belgium that on the occasion

of her witnessing, in a wood, just before a thunderstorm, a sudden

apparition of Salamanders in huge numbers, some among them

were distinguished by being marked with red instead of yellow.

In D’Orbigny’s ‘ Dictionnaire d’Histoire Naturelle’ (7, p. 307)

allusion is made to a specimen found near Bordeaux, which must

have been similar to the one described by Fr. v. Schweizerbath.

In Belgium, the markings are of a more or less bright yellow but

not orange, and in most cases they form interrupted stripes. In

Brittany the markings vary from sulphur- to chrome-yellow. Out

of 50 specimens obtained together, within a space of one hundred

square yards, last summer at Roscoff, about half had the stripes

uninterrupted but varying much in width. M. Ghidini, of the

Geneva Museum, having had occasion to examine 500 specimens

received alive from Stadtoldendorf in Brunswick, found that

about 400 had the two parallel stripes uninterrupted, or nearly

so, 50 had them much broken up, whilst the remaining 50 were

nearly entirely yellow, with the black reduced to spots or narrow

stripes. The specimens from the Harz and neighbouring hills, of

which I have seen many, vary in the colour of the markings from

chrome-yellow to a rather deep orange.

A male specimen from the Harz, in which the yellow colour

prevails, is represented on Plate XV.

Figures of the var. teniata are given by Wurfbain (338), Gesner

(18, ii. p. 80), Perrault (21, pl. 16. p. 77), Duvernoy (11, pl. xl.

fig. 1), Mme. Phisalix (22, pl. i.), and Durigen(10, p.577). The

descriptions of Leydig (19), Lataste (16),and Martin and Rollinat

(20) are also applicable to it, as well as the var. B of Dumeril and

Bibron (9) and the vars. f to & of Schreiber (27).

Five specimens are represented in text-fig. 102, to give some

idea of the variations in the markings :—

a, from the Harz, is exceptional in having the spots much

reduced in size; spots on the belly numerous, moderately

large and roundish.

b, from Maredsous, Belgium, represents the condition most

VARIATIONS OF THE SPOTTED SALAMANDER, ae

Var. teniata.

Tur SporteD SALAMANDER.

frequently met with in France, Belgium, and Germany ;

the ventral spots are confluent into a broad band on

each side.

c, from Roscoff, Brittany, is selected out of fifty specimens

as having the spots confluent into two stripes, and yet

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VARIATIONS OF THE SPOTTED SALAMANDER. 341

much reduced in width; the small amount of yellow on

the parotoids is also very exceptional; the belly is

entirely black.

d, also from Roscoft, is remarkable for the regularity of the

two yellow dorsal bands ; the yellow predominates on the

lower parts.

e is selected out of many from the Harz as showing the

preponderance of yellow, the black of the upper surface

being reduced to a few markings; the throat and _ belly

are yellow with scanty black spots.

Leaving out the specimen from Rome mentioned by Duméril

(possibly through some error of locality), the habitat of this variety

is restricted to France, Spain and Portugal, Belgium, S. Holland,

Germany, and Switzerland. On its occurrence alongside with the

typical form, see above, p. 333.

LV. Dr. KAMMERER ON THE VARIATIONS OF

SALAMANDRA MACULOSA.

In the introduction to this paper I have expressed surprise at the

distinction between the typical form and the striped form not

having been brought out more clearly by the authors who have

dealt with the varieties of this species.

This applies also to the most recent worker on the subject,

Dr. Kammerer (14, p. 69), some of whose highly interesting

observations are here reproduced. As he has accompanied them

with remarks on the correlation between the coloration and the

conditions under which the individuals occur and has drawn

provisional conclusions with which I cannot always concur, I

have appended my criticisms, inserted in square brackets.

It is possible, he thinks, to establish the existence of local modi-

fications as concerns the intensity of the yellow and its distribution

on the black ground-colour. Number, size, and intensity of the

yellow spots are in direct proportion to one another. One seldom

meets with specimens with few but large spots*.

[This statement is evidently meant to apply to the typical form

with isolated spots, but fails to express the state of things in cases

when several spots fuse together and are consequently large and

few ; the very yellow specimens, which are not so very unfrequent,

having the spots few in number. |

The contrary, many but small spots, occurs only in the var.

corsica ( fide Bedriaga)T

[This is perfectly true as regards Bedriaga’s specimen from

% “ Bs lassen sich namlich hinsichtlich der Sattigung des Gelb und der Vertheilung

desselben aut der schwarzen Grundfarbe lokale Abanderungen feststellen, die durch

eine Menge verschiedenartiger Faktoren beeinfiusst zu sein scheinen. Zahl, Grésse

und Sattigung der gelben Flecken stehen in direkter Proportionalitat zu einander:

man findet selten Individuen mit wenigen, aber grossen Flecken.”

+ “Das Umgekehrte, viele, aber kleine Flecken, tritt nur bei der var. corsica Savi

(vel. v. Bedriaga) auf.”

Proc, Zoou, Soc.—1911, No. XXIII. a4

342, MR. E. G. BOULENGER ON THE

Bocognano (now in the British Museum), but we must remember

that such a correlation is by no means constant in Corsica, for the

figure of the type of S. corsica, in Bonaparte’s ‘ Fauna Italica’

shows the number of spots not to be in excess of that of a typical

specimen from Italy, figured in the same work. ‘The spots on the

Bocognano specimen, although more numerous, are not smaller

than is usual in specimens from Hungary, Bosnia, Roumania, &c.,

or, for instance, the one from Italy, so beautifully figured by

Rusconi. It is also to be borne in mind that Savi in his original

description of jS. corsica ascribes to it fewer spots than to

S. maculosa. |

Where there is much yellow this is usually also strongly

intensified (dark straw- or orange-yellow), whilst scanty yellow is,

apart from rare exceptions, pale (pale sulphur or lemon) *.

[I cannot agree in the least with this statement, as out of over

50 specimens from the Harz Mts. which were recently received

at the Zoological Gardens, those in which the yellow constituted

the ground-colour were, as a rule, of a paler yellow than those in

which the black predominated. Again, in some specimens from

Dresden, as my father informs me, and in others from the

Harz, with the spots few and of small size, the colour was

decidedly orange, whilst in specimens from Brittany and Belgium

with much yellow, the latter varied from pale lemon to chrome.

A large number of specimens received alive from Hungary

(N. Becskeret), mostly with small, or very small spots, few in

number, had these orange, not yellow. |

The author then enumerates, with reference to his own

material and some indications in the literature, the local

differences in connection with the geographical distribution :—

1. In specimens from the hilly districts (Riva, Tyrol, 70-120

m., Schandau on Elbe, Saxony, 130 m.), and in those from the

North-German plain (Minden in Prussia, 50 m.) and from the

South of France and Spain (fide Bedriaga), the yellow is dis-

tributed in great profusion, the spots of the upper surface being

often confluent into broad longitudinal bands, or even becomes

the predominant colour, and in the 8. French and Spanish Sala-

manders the dark ground-colour disappears almost entirely.

Besides, the under surface of such specimens is strongly spotted

or even entirely yellow,

2. Inspecimens from Portugal (var. molleri Bedriaga) red spots

formed of a special pigment appear in addition to the mostly

numerous yellow spots, which here also invade the ground-colour.

3. The same remarkable appearance obtains likewise in many

specimens from the neighbourhood of Vienna (Hiitteldorf,

Moédling, Puckersdorf, Hadersdorf, Unter-Tullnerbach), in which

* “Wo viel Gelb vorhanden ist, da ist es gewohnlich auch stark gesattict

(Dunkelstroh oder Orangegelb), wahrend sparliches Gelb, von seltenen Ausnahmen

ahgesehen, blass (Lichtschwetel oder Citrongelb) aussieht. Bei einer massigen

Quantitat Gelb treten die Flecken bald in dunkleren, bald in helleren Schattirungen

auf, jedoch in der Regel nicht bei ein und demselben, sondern bei verschiedenen

Exemplaren.”

VARIATIONS OF THE SPOTTED SALAMANDER. 343

red of different shades (brownish red, greyish red, claret-red,

blood-red, vermilion, brick-red) is present, namely on the paro-

toids, on the forehead and vertex, round the eyes, at the angle of

the mouth, on the throat, and occasionally also over the whole

body. Otherwise the Vienna individuals are mostly provided

with moderately large and moderately numerous spots, the

intensity of the shade of which varies to the extent that in

different examples from one and the same locality they may be

pale or deep yellow.

4, Specimens from the Central Mountains of Germany, the

spurs and the lower region of the Alps (Salzburg, 450-638 m.,

Kaumberg, Lower Austria, 490 m., Meran, Tyrol, 350 m.,

Villach, Carinthia, 500-600 m., Jenbach, Tyrol, 530 m.,

Wochein Valley, Carniola, 510 m., Bozen, Tyrol, 250-300 m.,

Kapfenberg, Upper Styria, 500 m., Wiesing, Bavaria, 750 m.,

Mondsee, Upper Austria, 480 m., Kufstein, N. Tyrol, 490 m.)

show an average condition, 7.e., the mostly irregularly formed

and distributed spots on the upper surface being moderately large

and numerous, those on the lower surface being scanty and

pale; all sorts of shades of yellow are observable in different

individuals.

5. In specimens from the Alpine region, from the upper limits

of the vertical range of the species (Salzburg, 1000 m., Bozen,

Tyrol, 920 m., Appenzell, Switzerland, over 1000 m.), the black

ground-colour predominates, the yellow spots being small, few,

and very pale. The lower surface is usually unspotted. Accord-

ing to Walter Bendt, of Gratz, a specimen obtained on the

Schéckl Plateau at an altitude of about 1230 m. (highest altitude

on record) had only a few spots which were not of a well-defined

yellow, but more brownish.

6. Small number and size of spots are also shown by specimens

from S.E. Europe, viz., from Orsova (jfide Mojsisovics), Monte-

negro ( fide Werner), and Athens (fide Werner), and generally by

Turkish and Greek Salamanders, also by the specimens from the

extreme south of the distribution (Haifa, Syria, and var. algira).

Not everywhere are the local variations quite constant, viz.,

there occur in the Alpine region (Alpine valleys up to 600 m.,

de Werner) rather strongly and largely spotted specimens,

whilst in the neighbourhood of Vienna, among an average of

strongly spotted specimens, there are also some, if only a few,

with small and yellow spots. Indications of red spots on the

parotoids, on the inner edge of the upper eyelids, as well as

between the angle of the mouth and the lower border of the eye,

are also to be found in isolated specimens from Lower Styria and

the Bavarian Alps.

An absolute constancy with regard to the multiplicity of the

factors which influence the markings, many of which may, of

course, counteract one another in the same locality, cannot be

expected.

A precise test of the geological, climatic, and meteorological

23%

344 MR. E. G. BOULENGER ON THE

conditions of the localities of a great number of specimens has

shown that the colour-variations can be grouped ona geographical

basis * :—

1. A warm climate increases, a cold climate attenuates the

markings and their intensity.

[Salamanders from Belgium and the Harz district, with cold

climates, in the north of the distribution of the species, have on

an average large markings, the Harz specimens in. particular

being often bright yellow or orange, whilst specimens from

S.E. Europe, N. Italy, to say nothing of extreme southern speci-

mens (Syria and Algeria), to which Kammerer himself alludes,

have, as a general rule, small and few spots. Specimens from the

hilly parts of Belgium, from most French localities, either at sea-

level (Boulogne, Brittany, Bordeaux) or at a considerable altitude

(Haux-bonnes, Pyrenees, 750 m.), are absolutely identical in the

general style of markings and in their colour, thus showing that

climate has no bearing on this character. |

2. Localities with very damp air and soil, provided in addition

with a great number of water-courses, favour the number, size, and

intensity of the yellow spots, whilst conversely dryness and

scarcity of water-courses produce a decrease in these respects.

[One does not well conceive the Salamander flourishing under

other conditions than the first. It is difticult to believe that the

Salamanders with much yellow, such as we have from Bordeaux,

S. Italy, and Asia Minor, are from damper localities than those

with small spots from Austria, Hungary, and Roumania. What

I have said above of the Salamanders of France is against

Dr. Kammerer’s contention. | '

3. In localities where the sub-soil consists of schists, igneous

rock, and sandstone, Salamanders are as a rule more numerous,

larger, and more intensively yellow-spotted than in calcareous

hills, where they are not found in such abundance; possibly this

may bring us back to proposition 2, calcareous hills being always

* “Hine genaue Priifung der geologischen, klimatischen und meteorologischen

Verhaltnisse méglichst vieler mir aus eigener Erfahrung und aus Museen durch

Belegeexemplare, sowie aus der Litteratur durch Beschreibungen der betreffenden

Exemplare bekannt gewordenen Fundorte hat ergeben, dass die Farbenabanderungen

an Stelle der geographischen Gruppirung auch in folgender Weise geordnet werden

kounen:

1. Warmes Klima unterstiitzt, kaltes Klima unterdriickt die Fleckenzeichnung

und deren Sattigung.

2. Gegenden mit starker Luft- und Bodenfeuchtigkeit welche woméglich ausserdem

reich an kleinen Wasserlaufen sind, begtinstigen “Zahl, Grosse und Sattigung der

gelben Flecken, wogegen Trockenheit und Armuth an Gewassern dieselben zurtick-

treten lasst.

3. In Gegenden, wo Schiefer, Urgestein und Sandstein den Untergrund bilden,

sind die Salamander in der Regel zahlreicher, grosser und intensiver “gelb gefleckt

als im Kalkgebirge, wo sie iiberhaupt nicht so haufig zu finden sind. Méglicher

Weise lisst sich dies auf Punkt 2 zuriick fiihren, indem das Kalkeebirge stets

trockener und &rmer an Gewassern ist, als das aus den ubrigen genannten Gesteins-

arten sich zusammensetzende Gebirge.

4. Ein ganz besonderes Uberhandnehmen der gelben Farbe und deren Sattigung,

sowie das Auftreten von rothen Flecken, findet auf Lehmboden statt, wogegen aut

schwarzen Humus das Umgekehrte obw altet.’

VARIATIONS OF THE SPOTTED SALAMANDER. $45

dryer and poorer in water-courses than those formed of the other

mentioned rocks.

[I am not sufficiently acquainted with the formations on which

the specimens in the museum were obtained to fully discuss this

proposition, but I wish to observe, on the testimony of my father,

who has examined large numbers of specimens from Belgium,

found on Carboniferous and Devonian limestone, and from

Brittany, on granite, that Salamanders from these places agree

absolutely in the extent of the spots and in the comparatively

pale colour of the same as well as in size. This is true also of the

Salamanders found over the greater part of France (var. teniata,

with markings hardly ever orange), and yet the localities about

which we have definite information are on the most different

geological formations :-—a. Ambleteuse, near Boulogne (Kim-

meridge and Upper Oolite, clay with lime); 0, c. St. Malo and

Roscoff (Granite); d. Bordeaux (Pliocene and Oligocene clays) ;

e. Marly, near Paris (Oligocene limestone); /. Haux-bonnes,

Pyrenees (Cretaceous limestone). This list clearly indicates

that, in France at any rate, the subsoil has nothing to do with

the coloration of Salamanders. |

4, A quite special increase of yellow colour and its intensity,

as well as appearance of red spots, takes place on a clay soil,

while, on the other hand, the reverse takes place on black

humus.

Dr. Kammerer coneludes by observing that the above correla-

tions are subject to many irregularities, and that he would

formulate them with reserve, were it not a fact that these

irregularities or apparent exceptions are almost invariably to be

satisfactorily explained by the simultaneous occurrence of opposed

factors in one and the same locality (e. g., water poverty but clay

soil, or warm climate but calcareous hills, &c.) and that it is

therefore necessary to make a complete investigation of the

physical conditions of each separate locality.

[Suffice it to add once more that the large Belgian Salamander,

copiously marked with yellow, occurs in great abundance in places

offering a combination of the two principal factors which, ac-

cording to Kammerer, would produce small size and scarcity of

markings, viz., a dark soil rich in lime and a cold climate.

It is also important to observe that, contrary to Kammerer’s

results, our blackest specimen of the typical form is from a hot

locality (Ticino) at an altitude of only about 600 metres, our

yellowest from the highest altitude on record, 1600 metres

(Calabria). It might be objected that in the case of the latter the

altitude is compensated by the latitude, but that will not answer,

since at a more southern latitude, and almost at sea-level, on the

north coast of Africa, paucity of yellow markings is the rule. In

the var. teniata, a totally black specimen has been recorded from

the Siebengebirge (Rhenish Prussia), whilst the opposite extreme

of almost wholly yellow specimens is on record from as distant and

dissimilar localities as the Harz, Bilbao, Toulouse, and (?) Rome. ]

346 MR. HE. G. BOULENGER ON THE

Since the publication of the above conclusions, Dr. Kammerer

has instituted a series of experiments with the object of showing

that the colour and dampness of the soil has an influence on the

increase or reduction of the yellow markings, and the results of

some of the experiments have been recently published, with figures,

in the German paper ‘ Natur’ (15).

In view of the geographical distribution of the typical form and

the var. teniata, which I have endeavoured to trace from a large

material, Dr. Kammerer’s results are surprising, for the pictures

given by him show the offspring of a female of the typical form,

presumably from Austria, to belong to the var. teniata. Con-

sidering the enormous number of specimens of the two forms

which he has kept in his terrarium, is the possibility excluded of

some confusion having taken place? Or may not the male have

belonged to the latter variety, and have transmitted his characters

to the offspring? Then, again, as to the changes in markings

taking place in the same individual in the course of growth, may

not some error of identification have crept in? After the study

I have made of the disposition of the markings, I can hardly

refrain from expressing a doubt as to the middle specimen of the

left-hand lower series on fig. 1 being the same as the one to its

right, for it will be observed that the two yellow stripes or series

of spots are much more distant from each other in the younger

stage than in the older, and this is a change difficult to conceive

to have taken place in one and the same individual. In fact, were

it not for the statement of so highan authority as Dr. Kammerer,

whose experiments appear to have been conducted with so much

skill, care, and patience, 1 think I would not have hesitated

in pronouncing the two figures in question to have been taken

from different individuals.

REFERENCES.

1. Auprovanpr, U.—De Quadrupedibus Digitatis Oviparis.

Lib. i. (Bonn, 1637.)

2. Bepriaca, J. von.—Beitrage zur Kenntniss der Amphibien

und Reptilien der Fauna von Corsika. Arch. f. Naturg.

1883, p. 252.

3 — Die Lurchfauna Europa’s.—I1. Urodela. Bull. Soe.

Nat. Moscou, 1896 (1897).

4. Berra, E. de.—Monografia degli Anfibi Urodeli Italiani.

Mem. Istit. Veneto, xi. 1864, pp. 495, 516.

. Bonaparte, C. L.—Iconografia della Fauna Italica. Vol. 11.

. CamErano, L.—Monografia deel Anfibi Urodeli Italiani.

Mem. Acc. Tor. (2) xxxvi. 1884, p. 405, 2 pls.

. D’Orsieny, C.— Dictionnaire d’ Histoire Naturelle, xi. (Paris,

1849.) Art. Salamandre.

Doumercur, F.— Essai sur la Faune Erpétologique de

VOranie. (Oran, 1901.)

Dumirit & Bipron.— Erpétologie Générale, ix. (Paris, 1854.)

10.

pls

12.

13.

14.

15.

16.

18.

20.

al.

22.

23.

24.

25.

26.

27.

28.

29.

30.

31.

32.

33.

VARIATIONS OF THE SPOTTED SALAMANDER, 47

DériceN, 6.—Deutschlands Amphibien und Repiilien.

(Magdeburg, 1897.)

Duvernoy, G. L.—Planches du Réene Animal. Reptiles.

(Paris. )

Fung, A. F.—De Salamandrae Terrestris. (Berlin, 1827.)

Gusner, C.—Historia Animalium. Lib, ii, (Frankfort, 1617.)

Kammerer, P.—Beitrag zur Erkenntniss der Verwandts-

schaftsverhiltnisse von Salamandra atra und maculosa.

Arch. f. Entwickelungsmech. xvii. 1904, Heft 2.

Vererbung erzwungener Farb- und Fortpflanzungs-

verinderungen. Natur, vi. 1910, p. 94.

Lataste, F.—Kssai d’une Faune Herpétologique de la

Gironde. Actes Soe. Linn. Bordeaux, xxx. 1876.

LATREILLE, P, A.—Histoire Naturelle des Salamandres de

France. (Paris, 1800.)

Laurenti, J. N.—Specimen Medicum. (Vienna, 1768.)

Leypre, F.—Ueber die Molche (Salamandrina) der W iirttem-

bergischen Fauna. Arch. f. Naturg. 1868.

Martin, R., & Ronturat, R.—Vertébrés sauvages du Deé-

partement de l’Indre. (Paris, 1894.)

Perravuttr, C.—Descriptions anatomiques de deux Sala-

mandres. Mém. Ac. Se. Paris, 111. pt. 3, 1734, p. 77, pl. xvi.

PuisALix-Picor, Mme.—Recherches.... sur les glandes a ~

venin de la Salamandre terrestre. (Paris, 1900.)

Reber, J. E., & Haun, C. W.—Fauna Boica. (Nurenberg,

1832.)

RoéseL von Rosennor.—Historia Nat. Ranarum Nostratium.

(Nurenberg, 1758.)

Rusconr, M.—Histoire Naturelle, Développement et Méta-

morphose de la Salamandre terrestre. (Pavia, 1854.)

Savr, P.—Descerizione della Salamandra corsica. N. Giorn.

Lett. xxxvil. 1838, p. 208.

ScureiBeR, E.—Herpetologia Europea. (Brunswick, 1875.)

SCHWEIZERBATH, ExisE v.—Eine rote Farbenvarietiit von

Salamandra maculosa. Ber. Senck. Ges. 1906, p. 119,

pl.iv. Also Bl. f. Aq. u.Terr. Kunde, xx.1909, No. 24, p. 382.

SeoAne, V. L.—Identidad de Lacerta schreiberi (Bedr.), y

Lac. viridis vax. gadovii (Boul.) é investigaciones herpeto-

logicas de Galicia. (Corunna, 1884 [1885].)

Sturm, J.—Deutschlands Fauna. Part ii. Heft 2. (Nirn-

berg, 1799.)

Werner, F.— Untersuchungen tber die Zeichnung der Wir-

belthiere. Zool. Jahrb., Syst. vi. 1892, p. 155.

Die Reptilien und Amphibien Oesterreich-Ungarns

und der Occupationslinder. (Vienna, 1897.)

Wourreain, J. P,—Salamandrologia, (Nurenberg, 1683.)

EXPLANATION OF PLATE XV.

Salamandra maculosa.—Yorma typica, 8, from Lugano; var. molleri, 2,

from Portugal; var. teeniata, g, from the Harz.

348 MR. R. LYDEKKER ON

17. On the Mountain Nyala, Tragelaphus burton.”

By R. LyDEKKER.

{ Received December 1, 1910: Read February 21, 1911. ]

(Plate XVI.t and Text-figure 103.)

About the 10th of September I received a communication from

Mr. Rowland Ward to the effect that the skin, skull, and horns

of an apparently new Kudu-like Antelope had been received at his

establishment in Piccadilly from Mr. Ivor Buxton, by whom the

animal had been shot in Abyssinia. An inspection of the

specimen a few days later fully convinced me of the correctness

of Mr. Ward’s diagnosis ; and I accordingly wrote a letter to the

‘Times,’ which duly appeared in that journal on September 23rd,

1910, under the heading of “‘ A New African Antelope.” In that

letter it was stated that the specimen had been killed on the Arusi

plateau of Gallaland, in Southern Abyssinia, at an estimated

height of about 9000 feet above sea-level, and that it apparently

indicated a new species of Antelope in some degree intermediate

between the Nyala (7ragelaphus angast) and the Kudu (Strepsiceros

capensis), but rather nearer to the latter than tothe former. In

conclusion, it was urged that the specimen ought to find a

permanent home in the British Museum. It was also suggested

that the species might be known as the Spotted Kudu.

Shortly after the appearance of this letter, Mr. Buxton wrote

to say that he would be pleased to present the specimen to the

Museum, and likewise giving full and more precise information

with regard to the locality where it was obtained. The matter

having thus become public, I decided that the time had come to

give the animal a scientific name; and Strepsiceros buatoni was

accordingly proposed by myself in ‘ Nature,’ vol. 84, p. 397, 1910.

It was, however, added that it might be deemed advisable to

merge the genus Strepsiceros in Tragelaphus, in which event the

title of the new species would be Z'ragelaphus buatoni. As to

locality, Mr. Buxton, after reference to his note-book, informed

me that the type specimen of the new Antelope was obtained

during the summer of 1910, to the west of the Arusi plateau of

Gallaland, in the Sahatu Mountains, at an estimated elevation of

9000 feet ; these mountains being situated some distance to the

south-east of Lake Zwei(Zuay). The ground on which the animal

was killed—-as I gather from a photograph taken by Mr. M. C.

Allbright, who accompanied Mr. Buxton on his trip—is of an

open and stony nature, with scattered dwarf bushes and tussocks

of grass.

Soon after the publication of the aforesaid notice in ‘ Nature’

I learned that Mr. Buxton had brought home the head-skin

* Published by permission of the Trustees of the British Museum.

7 For explanation of the Plate see p. 353.

. Pivoo LOW PIXE.

H.Goodchild, del et lth. A.S Huth, imp.

THE MOUNTAIN NYALA.:

TRAGELAPHUS BUXTONI.

THE MOUNTAIN NYALA. 349

and skull of a second male of the same species, and also the skulls

and horns of two other, and apparently younger, individuals of

this sex—all being members of the herd from which the type

specimen was obtained. Nor is this all the available material ;

for Mr. Allbright had likewise shot a fine old buck out of the same

herd, of which the skull and skin were brought home, the head

being mounted and the body-skin preserved flat. All these

specimens, thanks to the courtesy of their respective owners, I

have had full opportunities of examining. It should be added

that, according to Mr. Buxton, the females of the new species are

coloured like the males and are devoid of horns.

The second head brought home by Mr. Buxton and the complete

skin belonging to his companion represent darker-coloured and

apparently older bucks than the type; and after seeing these

specimens I came to the conclusion that the Sahatu Antelope is

nearer to the Nyala than to the Kudus. This opinion is recorded

in the ‘Field’ of October 22nd, 1910 (vol. 116, p. 798), where I

definitely stated that the species ought to be knownas 7ragelaphus

Suxtont. Finally, in the recently published sixth edition of

Mr. Rowland Ward’s ‘ Records of Great Game’ I have suggested

the name Mountain Nyala as the popular designation of the

species.

That the type specimen, which is a male, and stands, as

mounted, 4 feet 44 inches at the withers, represents an Antelope

belonging to the Kudu and Bushbuck group is self-evident, and

further words on this subject are therefore superfluous. Kudus

and Bushbucks are admittedly near relatives; and the main

distinetion between the two is to be found in the characters of the

horns. In adult bucks of both species of Kudu the hind surface

of the basal portion of the horns is rounded and devoid of an

external ascending keel, although there is a well-marked front or

inner keel. The horns of Bushbucks, on the other hand, have a

pronounced keel at the back of the basal portion, in addition to

which is a distinct front keel; and, in consequence, the surface

between these two keels is flat, instead of round. This flattened

interearinal surface is, in fact, continued nearly to the summit

of the horn, forming a spiral round the axis. In adult Kudus a

second keel is developed higher up on the horns, and such a

flattened surface consequently exists only near the tip. In the

horns of a young Kudu, on the other hand, there isa second keel ;

and it follows that the horns of the adult may be compared to

Nyala horns with an additional basal portion devoid of a second

keel. This shows that the distinction between Strepsiceros

and Tragelaphus is very slight indeed.

Kudu-horns are, however, distinguished from those of the Bush-

buck group by their open, corkscrew-like spiral of three complete

turns, those of Bushbucks never having more than a couple of

such turns.

In Kudus the males and females are alike in colour, or nearly

so, the coloration on the body consisting of vertical white stripes,

350 MR. R. LYDEKKER ON

with a tendency to spotting on the hind part of the flanks of the

smaller species. Both species have a mane and dorsal crest, at

least in the males; but while the larger kind has none, the Lesser

Kudu carries one white gorget on the throat and another on the

chest. The tail is of moderate length and not very bushy.

In the Bushbuck group the females are frequently—although

by no means invariably—brighter coloured and more profusely

striped and spotted than the males; the markings on the body,

when fully developed, comprising both stripes and spots. The

males, at any rate, have a dorsal crest; and there are often two

gorgets on the throat and chest, although these are reduced to one

in the Nyala. The tail is in most cases relatively long and more

bushy than in Kudus.

The conformation of the horns affiliates Mr. Buxton’s Antelope

to the Bushbuck group (Vragelaphus*), from all the other

members of which it is distinguished by its superior size. From

the Nyala, which makes the nearest approach in this respect,

Tragelaphus buxtowi differs in the (reported) identity of the colour

in the two sexes, in the shorter coat and less bushy tail of the

male, the presence of two white gorgets on the throat and chest,

the absence of any marked difference in the general colour of the

lower part of the legs from that of the body, and also in the more

open spiral formed by the more massive horns.

In the type specimen (Pl. X VI.), which, as already mentioned,

is an approximately full-grown but young buck, the horns form

about one complete turn, and have the general characters of those

of the Nyala, although relatively heavier, and diverging much

more outwardly, with a very open spiral. They are obliquely

ridged at the base, and the long smooth terminal portion is worn

yellow at the tip. The length along the outer curve is 37 inches,

the basal girth 97 inches, and the tip-to-tip interval 21 inches.

The coat is rather long and coarse, its general colour being

speckled brown-fawn, passing into dull tan on the sides of the

face, and becoming darker on the front surface of the muzzle, and

chocolate-brown on the forehead above the white chevron, which

is not very conspicuous. The under- parts are lighter, but on the

front of the fore-legs and the lower part of the hind pair the tuft

becomes considerably darker. There is a short dark brown mane

on the neck, continued backwards as a mingled brown and white

dorsal crest. The bushy tail is white beneath. The ears, which

are much of the same type as those of the Nyala, are of moderate

width, bluntly pointed at the tip, and tubular for a considerable

distance at the base; most of the long hairs on the inner edges

being white, as is also much of the inner surface of the outer

margin.

The white markings include a not very distinct chevron between

the eyes, the usual patches on the sides of the muzzle and chin, a

pair of spots on each side of the face below the eye, and a smaller

* Limnotragus \ regard as a subgenus.

HE MOUNTAIN NYALA. Sot

Text-fig. 103.

Head of an old Buck of the Mountain Nyala belonging to Mr. Allbright, from a

photograph supplied by Mr. Rowland Ward.

352 ON THE MOUNTAIN NYALA.,

and fainter one behind the same, a narrow but deep gorget on the

throat, and a much wider but less deep one of a more lunate shape

on the upper part of the chest. A curved row of nine spots—

some of which are much fainter than the rest—extends from a

point about over the head of the thigh-bone to the back of the lower

part of the shoulder. There is another white spot on each side of

the buttocks. The inner surface of the thighs and of the upper

portion of the fore-legs is dirty white. A white area occupies

the back of each fore-leg below the knee, extending on to the outer

and inner surfaces of the limb, but not reaching the pastern ;

and a somewhat similar area occurs on the hind-leg, extending

slightly above the hock. There is a pair of white oval spots on

each fetlock some distance above the hoof.

The second head obtained by Mr. Buxton is that of a much

older buck, as is attested by the horns, which have closely approxi-

mated rings at the base, are of greater length, and are much

battered on the front surface, and worn away at the tips, of

which the left one is broken. They form about one turn and a

quarter, and have a more upright direction than in the type

specimen; in both of which respects they are more Nyala-like.

Although the buck to which this head belonged was a member of

the same herd as the type, the coat is considerably longer and

looser, especially on the throat, where it forms an incipient fringe.

The colour is also darker and greyer, being a greyish brown,

comparable to that of a Waterbuck. The face is likewise distinctly

darker, the whole of the lower portion being of a chocolate-brown

like that of the forehead, and the tan restricted to the area round

the eye, behind which isa small white patch. There is a tendency

to rufous in the hair round the muzzle, and also in that between

the horns. In consequence of the darker colour of the rest of the

face, the white frontal chevron is much more conspicuous than in

the type specimen. The upper throat-patch or gorget is also very

conspicuous, and is continued by means of scattered white hairs

almost to the lower gorget.

The mounted head belonging to Mr. Allbright (text-fig. 103) is

likewise that of an old buck, and agrees in essential characters

with the one last-mentioned. ‘The body-skin of the same animal

differs from that of the type not only in its longer and darker

hair and the greater development of the dorsal crest, but likewise

in the presence of two indistinct vertical white stripes—one con-

siderably longer than the other—on the hind-quarters, with faint

traces of a still shorter third one. In this respect the specimen

makes a further approximation to the Nyala.

The skull is not yet cleaned, but does not apparently present

any very distinctive features of generic importance. It is true

that the base of the horn-cores lacks the pronounced front keel

found in the Kudu, but as this keel is also lacking in the Lesser

Kudu, it cannot be regarded as a feature of much importance.

On the whole, the Mountain Nyala seems to come nearest to

the species from which it takes its name, although in the general

;

ON SOME GIBBONS RECENTLY LIVING IN THE GARDENS. 353

form of the head and the character of the tail it is distinctly

Kudu-like.

In conclusion, I may reiterate my opinion that Zragelaphus

buetoni tends to connect the Bushbuck group so closely with the

Kudus as to render the generic separation of the latter from

Tragelaphus (in which, as already stated, I include Limnotragus

as a subgenus) inadvisable.

EXPLANATION OF PLATE XVI.

The type specimen, a subadult buck, of the Mountain Nyala,

Tragelaphus buxtoni.

18. Observations on different Gibbons of the Genus

Hylobates now or recently living in the Society’s Gardens,

and on Symphalangus syndactylus, with Notes on Skins

in the Natural History Museum, 8. Kensington. By

Dr. F. D. Wetcx, F.Z.S.

[Received November ‘29, 1910: Read February 21, 1911.]

Very few specimens of the genus Hylobates arrive in good health

in England or live for more than a few months, but during the

last five years the Society has exhibited examples of different

species, of which the following are now living in the Gardens :—

A male of the rare H. hainanus from Hainan, which arrived in

December, 1907; a male H. Jewciscus from Borneo, which arrived

in May 1908; andafemale 4. agilis from Sumatra, which arrived

in December 1905. In addition to these an adult male

Symphalangus syndactylus in good health arrived on October 26,

1910, and is the finest specimen the Society has yet received.

As some external characters and the coloration and voice can be

studied only in living specimens, some observations I have made

on these genera may be worth recording.

In the genus Hylobates, from observations taken at intervals on

four males living in the Gardens, I am strongly of the opinion

that the development of certain parts of the external genital

organs is delayed to a later period of life than is the case in

Anthropopithecus and Man, and, so far as I can discover, no notes

have been published on this subject. My attention was first

drawn to the subject by Mansbridge, keeper of the Society’s Apes,

but I have not accepted any information from him until I have

confirmed it myself, and i am well aware that several mistakes

have previously been made about the sex of Hylobates on account

of the large clitoris being mistaken for the penis (see Pocock,

P. Z.S. 1905, vol. ii. p. 169). I have made a careful examination

of ail the specimens, and am certain that the sex isas I have

stated.

The male H. hainanus has altered greatly in the external

354. DR. F. D. WELCH ON SOME GIBBONS

genital organs. On arrival in December 1907 it was in good

health and measured about 14 inches from crown to ischial

callosities, and there were then no signs of a scrotal bag, the skin

between penisand ischial callosities being quite flat. About the end

of March 1909 a scrotal bag gradually formed, and by the first

week in July 1909 this could be seen quite easily when the animal

was at the opposite side of the cage with its hind legs separated.

The scrotal bag had then reached its present size and appearance,

being just over an inch long with a broad base and tapering

rapidly towards a point, thickly covered all over with short black

hair, and placed well above and in front of the anterior ends of the

ischial callosities, there being a space over half an inch between the

eallosities and its posterior surface. The scrotal bag is not at all

pendulous, and the long axis runs downwards and forwards. At

that time, July 1909, the animal’s height, measured against the

bars when sitting, was 16 inches. It was impossible to measure

this male out of the cage or to handle it, and as all Hylobates

specimens stoop when sitting, the real height of the animal when

hanging at full length was probably about 1 inch more than this.

- The measurement against the bars was repeatedly taken to ensure

accuracy. Only one male H. hainanus has been exhibited in the

Gardens before, a younger animal, to the skin of which, now in

the Natural History Museum, I shall refer later.

Retention of the testicles, either within the abdominal cavity or

in the inguinal canal, is a common human abnormality. One or

both testicles may be thus retained, and in some cases they descend,

later in life, into the scrotal bag. I certainly do not think that this

male H. hainanus was abnormal, but that the absence of a scrotal

bag before March 1909 was the normal condition in the genus

and species, and my reasons for this opinion are as follows :—

The male H. lewciscus from Borneo, now in the Gardens, had

on arrival no scrotal bag, and in December 1909 I madea careful

digital examination of the external genital organs. The skin

between the penis and ischial callosities was slightly wrinkled, but

would not stretch when pulled, and did not hang down when the

animal was standing erect. There was nothing to be felt under

the skin between penis and callosities which I could say with

perfect certainty were the testicles, and consequently these organs

must be exceedingly small in proportion to the size of the animal.

This H. lewciscus was in good health on arrival and has continued

so, andat the present time (November 1910) there is no sign of a

scrotal bag. The animal at present measures 14 inches from

crown to callosities when sitting. Exactly the same condition

of no scrotal bag or testicles to be easily felt was found in two

other males I examined, one 14 inches high from crown to

callosities, the other 11 inches. This condition in these four Hylo-

bates males was very different from that of a young Anthropo-

pithecus troglodytes | examined, as in this latter male the scrotal

bag was large and well developed, being seen quite easily at a

distance of fifteen feet, and both testicles could be felt easily.

‘|

Mes

ee ey ST TES

Sa SS aU al ges ER gd a Ciesla we oh

ee a

RECENTLY LIVING IN THE GARDENS. 305

Tf is tomy mind most unlikely that four Zylobates males should

arrive in the Gardens one after the other with abnormal external

genital organs, and I think that in these cases the absence of

scrotal bag and small size of the testicles during the early years of

life were normal, and this opinion is strengthened by the condition

of some young skins in the Natural History Museum, which I

examined by kind permission of Mr. Oldfield Thomas. The skins

in that collection are of different ages, sizes, and species

(1. hainanus, H. hoolock, H. lar, H. leuciscus, H. agilis; FH, agilis

martiniand pileata, H. miilleri, H. leucogenys, H. gabrielle). Some

of these skins, those of old males, have a large and conspicuous

scrotal bag; in others, younger males, the scrotal bag is only

beginning to form and not so obvious, while in some of the

youngest skins, one of which is the male 1. hainanus above men-

tioned, it cannot be seen or felt. In these the skin between penis

and ischial callosities is perfect and has not been cut at all, so it

is quite certain no scrotal bag existed during life.

The delayed development of the scrotal bag and diminutive size

of the testicles correspond to the late commencement of menstru-

ation, and on this latter subject few observations have been made

in Hylobates.

I have compared the size of the skull as felt through the skin,

and also the canine and other teeth, of the female 1. agilis now

living in the Gardens with skulls in the Museum which are

obviously adult, and this comparison shows that the Society’s female

H. agilis is almost adult at the present time; although it has

been in perfect health since arrival in December 1905, it has not

yet begun to menstruate. Mr. Pocock has also recorded that the

female H. hainanus previously exhibited did not begin to

menstruate until almost adult (P. Z. 8S. 1905, vol. ii. p. 169), and

from these two cases it seems to be the normal course that in

Hylobates menstruation is delayed until a much later period than

is normal in Man.

When the female H. hainanus just mentioned was living in the

Gardens, I noticed that it was in proportion to height more

slenderly built in both body and limbs than other species (such as

H. agilis, H. lar, H. leweiseus, and H. hoolock) then or recently

living. As the animal had one fore limb, which it rarely used,

partially paralysed, I thought at that time the slender build was

the result of poor health and not a specific character. This being

the first female brought either alive or in skin to England, so far

as was known, there was nothing to compare it with, but since

then the male now in the Gardens arrived and showed the same

slender body and limbs. As this male has been in perfect health

since arrival in December 1907, and as its appetite is good and

it has not become stouter at all, although it has grown taller,

I think it is quite justifiable to state from these two living

specimens that I. hainanws is more slenderly built in body and

limbs than other species such as H. agilis, H. lar, H. hoolock,

and ZH. leuciscus.

356 DR. F. D. WELCH ON SOME GIBBONS

A comparison of the external genital organs of the male

H. hainanus with those of Symphalangus syndactylus shows

certain differences. In H. hainanus the distal half of the penis,

which is covered by mucous membrane, is a dull red, making

that organ very conspicuous in the otherwise jet-black coloration.

In S. syndactylus the penis is black all over, and both it and

scrotal bag are very small indeed in proportion to the size of the

animal. When hanging the penis is on a level with the ischial

callosities, and not above themas in H. hainanus. In S. syndac-

tylus the large tuft of long hair growing from below the penis and

between the callosities, and spreading out as it runs downwards and

backwards between the animal’s legs, is most conspicuous, hiding

a large part of the ischial callosities from view when the animal

is hanging and seen from below. In the skims of Hylobates in the

Museum the hair on the scrotal bag in old malesis no longer than

that on other parts of the body, except in one very aged H. hoolock,

and in this single specimen it is not nearly so long in proportion as

in S. syndactylus ; in skins of S. syndactylus males it is very long

indeed, as in our living male.

I might add that in H. hoolock and H. leuciscus the penis is

black all over, even in the distal half covered by mucous membrane.

H. hainanus presents three points distinct from H. hoolock as

shown in living specimens.

When Mr. O. Thomas described the type of H. hainanus (Ann.

Nat. Hist. ser. 6, vol. ix. p. 146), he was doubtful as to the

validity of the species of Hylobates; Mr. Pocock (P. 4. 8. 1905,

vol. ii. p. 169), from examination of a living female example of

H. hainanus, supported the distinctness of the species, and my

own observations on our living male confirm this.

The differences between living specimens of H. hainanus and

H. hoolock are as follows :—

1) H. hainanus is more slenderly built in body and limbs

than H. hoolock, as I have already pointed out, even when in

perfect health.

(2) In living males the colour of the penis is different. In

H. hainanus the proximal half covered by the skin is black,

and the distal half covered by mucous membrane is dull red. In

H. hoolock, on the other hand, both proximal and distal halves are

black.

(3) In two living specimens of H. hoolock, male and female, the

hair on the crown of the head lay quite smooth and there was no

erect crest whatever. In the male H. hainanus now alive the

hair on the crown of the head stands erect in a crest, as it

did in the female, as Mr. Pocock remarks (P. Z. 8. 1905, vol. ii,

elisa)

: I Eat add that the hair on the crowns of the living specimens

of H. agilis and H. leuciscus is quite flat and there is no erect

crest whatever. H. hainanus is the only species that I have yet

seen which has an erect crest on the crown; it also has a short

beard on the chin.

RECENTLY LIVING IN THE GARDENS. Son

There are two points in ZH. agilis worth mentioning which are

well shown in the female now living :—

(1) The tongue is of a dark bluish purple, the colour when

first seen suggesting that the organ had been very severely bruised

and that there was much extravasated blood in it; the absence

of any swelling, however, soon showed it to be quite normal. In

the other species (7. hainanus, H. leuciscus, and H. lai) the tongue

is, as one would expect in a genus so near to Man, of a pale reddish

colour, as is also the case in Symphalangus syndactylus.

There is considerable resemblance between young specimens of

H, agilis and H. lewciseus, and consequently this dark bluish-

purple tongue of 7Z. agilis should be of value in distinguishing the

species from HH. leuciscus, in which, as already mentioned, the

tongue is pale red. So far as I can discover, this peculiarly

coloured tongue of H. agilis has not been previously noted.

(2) In addition to the common chatter of excitement and

laugh which all species of Zylobates make, H. agilis has a loud ery,

quite distinct from the “ hoo hoo hoo” &e. of H. hainanus and the

“hah hoo hah hoo” &e. of H. hoolock (see Pocock on H. hainanus,

P. Z. 8. 1905, vol. ii. p. 176), and I have not as yet heard either

HT. lar or H, leuciscus-utter a similar sound. This peculiar ery of

H. agilis is best described by a word “ whopp,” and is shouted

out, the mouth being wide open and the throat conspicuously

dilated. It is only uttered once and then silence ensues for

about thirty seconds, then another “‘whopp,” then a period of

silence, then another “ whopp,” and so on. Usually this cry

continues for about ten minutes, but a few times I have heard

it continued for over an hour ; and very occasionally the ‘ whopp”

is prolonged into a long loud screech lasting a minute or longer,

the mouth being kept wide open the whole length of the screech

and the throat dilated. I have never heard it uttered when

playing or quarrelling with the H. hainanus (both species utter

the common chatter at these times), but only when the H. agilis

is swinging by itself. It is also a quite different sound from

anything uttered by S. syndactylus.

The voice of Symphalangus syndactylus is remarkable for the

variety of cries, which follow rapidly one after another, and at

least five sounds can be distinguished, with all of which the

gular bag dilates. Anyone with their eyes shut would certainly

think there was more than one animal in the cage. These cries

are :—

(1) The usual and loudest, best described by the words “ woe,

woe, woe,” &c., repeated rapidly and for a variable number of

times, sometimes forty or fifty. Judging from the deafening noise

the animal made with the “ woe woe” &e. on the first occasion I

heard it, I should think it could be heard much further off than

an adult H. hoolock, but unless one hears two adults shouting

against one anovher, it is very difficult indeed to be certain which

of the two has the louder ery.

(2) A gurgling noise, best described by the word ‘“ moo,” drawn

Proc. Zoou. Soc.—1911, No. XXIV. 24

358 MR. E. GC. OBERHOLLZER ON

out over several seconds and sounding somewhat like a human

being beginning to vomit. It follows rapidly after the ‘“ woe,

woe, woe,” &e., as a rule, and the gular bag dilates to the greatest

size with this sound, occasionally reaching a diameter of nearly

8 inches. The ‘‘ moo” is made mostly during inspiration.

(3) What I can best describe as a wailing-shriek like the word

‘“‘wair” shrieked out for twenty or thirty seconds and sometimes

longer, the voice being alternately raised and loweredalittle. It

usually follows rapidly after the ‘‘ woe, woe, woe,” &c., and 1s about

as often heard as no. 2—the gurgling “moo.” It is, however,

louder than the ‘‘ moo.”

(4) A “ho, ho, ho,” &e., repeated, as a rule, four or five times.

J have only heard it on a very few occasions and it 1s not nearly

so loud as the previous three sounds.

(5) A squeal somewhat resembling the noise made by some

Eagles. This is as rare as the “ ho, ho, ho,” &c., and not so loud

as the first three sounds.

When at rest the gular bag is black in sunlight and slightly

wrinkled, but on dilatation it becomes dull red. In its walk

S. syndactylus is bipedal like Hylobates.

EXHIBITIONS AND NOTICES.

March 7th, 1911.

Dr. A. Smirx Woopwarp, F.R.8., Vice-President,

in the Chair.

Tue Secretary exhibited a series of lantern-slides prepared

from photographs kindly given to him by Mr. Cart HAaGEenBeck,

Silver Medallist of the Society, and illustrating some of the most

remarkable features of Mr. Hagenbeck’s new Tierpark at Stellingen

- near Hamburg.

Mr. R. I. Pocock, F.L.8., F.Z.S., on behalf of Mr. Ernest

C. OBERHOLTZER, exhibited a large number of lantern-slides and

photographs illustrating the habits of Moose. Mr. Oberholtzer

had presented to the Society the series of enlargements exhibited

and communicated the following account of

Some Observations on Moose.

Of late it has become almost a heresy to associate big game

with any land but Africa. Like the bonneted old lady in the

corner, even America, humiliated, lives in the past; the stride of

the elephant steadily lengthens ; and one would suppose that all

save the jungle folk must soon be content with their barn-yards

THE HABITS OF MOOSE. 359

and menageries. What a consolation, therefore, to find in the

temperate zone a region where large wild animals are not only

numerous but increasing! Such was my own good fortune in

the forested lakeland tributary to Rainy Lake, Ontario, where

I spent five months in 1909 and the month of June 1910,

canoeing with a Red Indian. Besides numerous lesser animals,

we saw of that largest Deer—the Moose—nearly 500; 275 of

them in a single fortnight during the fly-season of 1910.

The Indians of this region all agree in saying that thirty years

ago the Moose was a rare, if not almost unknown, inhabitant.

Yet there are reasons to believe that at some remote time his

dominion was not unlike the present. On many of the rocks, for

instance, there are half-obliterated paintings attributed by the

Indians to a medicine man named Amo, who lived vaguely

“t’ousand years ’go”; and nearly all these paintings include the

figure of a moose. The horns and the hump on the back are

unmistakable.

The increase of the moose in the last thirty years has coincided

with the decline of the Indians; the natives still surviving are

generally a feeble lot, whose hunting-grounds are no longer far

afield. From his worst enemy, therefore, the moose has had

little to fear. Moreover, he has been almost without rival, for

the woodland Caribou that once roamed all over the region has

gradually withdrawn until to-day in a whole winter only an:

isolated band or so may be seen.

Another significant change in the region is the disappearance

of wild rice (Zizania aquatica), called by the Indians “ manomin.”

Formerly it was one of the Indians’ chief foods ; they gathered it in

great quantities every autumn and it attracted hosts of ducks.

To-day it is very rare ; in three thousand miles of canoeing I saw

it above water only two or three times and then always in small

patches. The explanation, I think, is connected with the moose.

Though writers never mention wild rice as one of his foods, there

can be no doubt that in these parts at least it is a favourite.

I remember one day we saw three moose feeding in a shallow

bend of the river. When we had frightened them away, I said

to my Indian: ‘ Billy, there are no lily-pads here. What were

they eating?” For reply, Billy thrust his arm under water and

pulled up a bunch of light green grass—the same that I had seen

hanging from the mooses’ mouths. It was wild rice.

Wherever in the Rainy Lake District moose are seen in large

numbers, careful investigation, I think, will reveal more or less

of this same wild rice, half-developed under water. The moose,

though they occur in all parts of the region, tend to congregate

in these special feeding-grounds. I have seen ten together,

seven of them bulls, in one bend of the Big Turtle River; and

every one was feeding on wild rice.

Browsing in the river, the moose usually walks out to the

height of his belly. If he feels no fear, he dips his head regularly

for a period of from ten to thirty seconds and lifts it during one

24*

360 MR. E. C. OBERHOLTZER ON

to fifteen seconds; such, at least, were my conclusions, after

roughly timing a number of feeding bulls. Not having a stop-

watch, I arranged with Billy to signal me every time the animal

either raised or lowered his head.

To get rid of flies, the moose often completely immerses

himself. I have seen large bulls swim out beyond their depth,

sink their heads, and be gone from sight a second or so, the

hump reappearing first. The bulls, as a rule, venture out much

farther than the cows; ten times to one where a moose 1s

overtaken in deep water by canoe it is a bull. This is partly,

I think, because the bulls have no responsibility in protecting the

calves, and partly because they seem to be much more afflicted

with flies than either cows or calves. Throughout June, July,

and often the greater part of August, the head and withers are

plastered with flies—particularly with little grass-green wedge-

shaped flies. When the bull immerses himself they merely hang

above the water till he reappears. Severai times when we have

overtaken a bull in mid-lake these flies have transferred them-

selves to our backs and there they stuck, stubborn to all slapping,

till we smoked them off over the fire. The cows and calves, on

the other hand, seem to be comparatively free from these pests ;

and I have often wondered whether the heat and blood ina bull’s

new antlers may not help to make him a target.

Last spring opposite our camp there was a bay, where we could

see the moose come to feed morning, noon, and evening. They

usually remained an hour or so, and often at night, too, we could

hear them splashing and grunting. In general, however, though

I have seen more moose at sundown than at any other tinie of the

day, I have not found them confining their visits to any particular

hours. If unmolested, many of the bulls during the fly-season

seem to remain in the water the greater part of the day; even

when disturbed they often return quickly. Nor do moose

cease to enter the water when the flies are gone. I have seen

a cow up to her belly on the first of November, a few days

before the river froze. They have a fondness for tramping

through the creamy grey ooze that covers the bottom of many

streams. Often you will see it sticking to their legs when they

stand on the shore; and I remember certain shallow lakes where

the bottom was channelled in all directions with their furrows.

Then, too, especially early and late in the year, they sometimes

take long swims. Last spring I saw two crossing the lake at a

place not less than two miles wide. In this way they often

become an easy prey to pot-hunters; one of my own canoemen

had thus slain a bull with an axe. Men have even been known

to jump on the backs of swimming moose.

Of the extraordinary tameness of moose in regions where they

are seldom molested there are many records. ‘They seem to have

a primitive curiosity about Man comparable to our own about

them; and this is especially evident in the untravelled Rainy

Lake District. Again and again we approached within fifty feet

THE HABITS OF MOOSE, 361

without causing uneasiness; and four times we came as near as

thirty feet in plain sight and took a dozen or more photographs.

I even changed my roll of films.

The most singular experience occurred on the 2nd of October,

1909. It wasabout half-past four o’clock in the afternoon ; the sun

was low, and we were looking fora place to camp. All at once

we heard the throaty grunt of a bull moose. Paddling quietly into

a shady bay, what was our surprise to see a little yearling cow

run out from the birch trees and come towards us whinnying !

The*next moment there was a loud crashing, and a tall heavy

two-year-old bull broke through the trees with a rush. I tooka

photograph at once, for I thought they would turn and flee.

Instead, as we drifted nearer and nearer, the little cow walked

out toward us along the edge of the shore as if for protection.

The bull, seeing us, checked his pace and waded stiff-legged into

the water as high as his knees. In evident fear of him, the little

cow kept whining and edging off, while he, advancing within fifteen

feet of us, lowered his head, turned broadside with one eye on the

cow, and sniffed at us suspiciously like a dog. The cow, as if

satisfied with her temporary immunity, now began to feed. The

bull made no attempt either to leave or to attack us. It was only

at last when I spoke, after thirteen pictures, that they took

fright ; and then the cow, instead of running away with the bull,

entered the water behind us and swam across to the other side.

The young moose, as a rule, seem to be less cautious than the

old. On the 11th of June, 1910, we paddled up the river to the

same bend, where the day before we had seen ten moose. This

day there were seven, and one by one they walked slowly away

into the woods, all except a little fellow that I mistook at first for

a cow. When we glided nearer, I saw two round knobs of horns

just peeping out of the forehead between his eyes and ears.

They were his first antlers. As we approached, he fidgeted a

little and looked at us with a mild sort of curiosity. Then down

went his head—but only halfway, for the tips of the ears still

protruded. Presently, completely reassured, he disappeared under

water. He was down on his knees. When he rose, he shook his

head till the long ears flapped against his cheeks. Then, whining

softly and stretching his neck forward, he took a step toward us.

I could almost have touched him with my hand ; but Billy, always

cautious, began to paddle backwards. Several times the little

fellow tried again to approach us—for exactly what purpose

I have never known. He seemed to be entirely friendly. At

last I spoke, and away he went.

There is a theory among some sportsmen who have only shot

moose that drooped ears are a sure indication of a wound. On

the contrary, the moose lowers his ears for as many reasons as any

other animal. Perhaps it is safe to infer that a moose with ears

always drooped is injured; but the pictures of the friendly little

bull show him with ears one moment erect, the next drooped, and

again out straight. Sometimes they are lowered on account of

362 MR. E. C. OBERHOLTZER ON

flies, sometimes to let the water run out, and almost invariably

when the moose is either apprehensive or at bay. A cow with

twin calves, which I photographed last spring, lowered her ears

when she began to feel uneasy about our presence. We were

only thirty feet away. At first she seemed not to mind; but

the repeated click of the shutter alarmed her. She appeared

to be wavering between friendliness and defiance. And I have

observed the same behaviour repeatedly under similar conditions.

Indeed, the moose seems to be an animal of mixed emotions.

Unlike the White-tailed Deer, he seldom bounds away at first

sight of man. He will havea good second look unless there is im-

mediate danger. Curiosity or indolence, or even friendliness (and

IT really believe in the last motive), frequently holds him until he

has had proof of evil intentions. Often, having crept up stealthily

upon a moose either by canoe or through the bush, I have been

amused to find that he was watching me wide-eyed all the time.

How often too, while the canoe passed, I have seen them peering

out unalarmed from behind their ambush. And certainly the

more they see of men without suffering injury, the more careless

they become. Passing up and down the same eight-mile stretch

of river at least once a day, last June, I was surprised at the

change that occurred in a week. At first I could never get

nearer than a hundred yards. Wealways went quietly and with as

few motions as possible, and we never pursued an animal that had

taken fright. At the end of the week hardly one of them would

budge until we were within seventy-five feet ; and if we returned

half an hour later, there they were again in the water. Some of

them never left, unless we turned toward them.

When we frightened them, it was either by unusual noises,

sudden movements, the scent of our bodies, or above all by the

sound of the human voice. No doubt, experience of the gun

changes the order; the scent then becomes of first importance.

I believe, though, that there is nothing alarming in the scent itself

except as it 1s novel or as it recalls deadly memories. The cow

and calves of which I spoke above may not have scented us, for

there was a slight breeze in our favour. With the young bull in

the water and the other bull and cow calf it was different. In

the former case the breeze blew straight away from us; in the

latter there was no perceptible breeze at all, but it would be idle

to question the powers of a sniffing bull at from 15to 20 feet. Itis

only the human voice, so far as I can discover, that never fails

the first time to cause a panic.. Whenever otherwise we were

admitted to the closest intimacy, it broke the spell at the first

word. There seems to be something unearthly about it that

jangles the moose to the very heart.

A peculiar feature of the moose is the bell. Mr. KE. T. Seton,

in his memorable ‘ Life Histories of Northern Animals,’ cites a

bell 18 inches long as unusual. For an average of all moose,

young and old, it certainly is; but, without actual measurements,

I have a strong impression nevertheless that for bulls of three,

THE HABITS OF MOOSE. 363

four, and five years even two feet is not rare. In cows and

yearling bulls the bell is usually small and delicate. In bills a

few years older I have often seen it swinging like a tassel. It

is the old bulls with large fully-developed antlers—the bulls

usually sought by sportsmen—that have no more than a thick

stump of a bell. The Indians believe, as Mr. F. C. Selous suggests

in his ‘ Recent Hunting Trips in North America,’ that it gets

torn off or dries up. Perhaps both may happen, or it may even

freeze off during the winter.

Except some of the yearlings and an old mother or so with

calves, the moose in the Rainy Lake District are all fat and sleek.

in September they fairly bulge. They seem lazy and satisfied, as

if they had found the true land of plenty. ‘They are not too

indolent, however, for play—at least not the younger ones. Once,

in June, I saw two of them (young bulls, I think, though I was

too far off to be sure) frolicking on a sand beach. They cantered

and reared and stopped short and wheeled round exactly like

young horses, and at last, without warning, bolted into the bushes,

Again, in September I saw a half-grown grey calf cavorting on

the edge of a bay. A very fat cow and a huge glossy bull in full

armour stood ankle-deep in the water, gazing at each other in

stupid indecision. They seemed to be equally infatuated. The

little calf meantime enjoyed all manner of caprices, just as if he

considered his mother’s new company a stimulating adventure for

himself. It reminded me of an exuberant boy tossing his hat in

the air.

During the rutting-season also we once tried the effect: of the

much-discussed “ calling ”—an art almost unknown in this region.

It was a warm afternoon—about three o'clock on the 30th of

September. We had just paddled a hundred yards or so beyond a

portage, when we heard the low, languorous, lustful, two-noted

grunt of a bull. In spite of the sun in our eyes, we thought we

saw him on the shore about three hundred yards to the right of us.

My Indian with unaided voice gave what seemed to me a perfect

imitation. Immediately, we heard the moose reply and wade out

into the water. The Indian repeated the call several times, and

the bull began to swim leisurely toward us. The call sounded so

easy that I could not resist trying myself. This seemed to make

no difference. The moose swam straight toward us. We then

kept still. When he was about one hundred feet away (a three-

year old bull, as we judged), he suddenly caught sight of us. With

evident surprise he turned and swam swiftly for shore. Whatever

else he had expected, I am sure he had had no thought of man—

even though my own call, at least, must have been seriously

defective. I should conclude, therefore, with Mr. Selous that,

where moose are unaccustomed to being deceived, a bull in a state

of frenzy may be attracted sometimes by other sounds than the

perfect call.

The moose, like all creatures, sometimes makes fatal miscalcu-

lations. A real tragedy was recently reported to me ina letter from

364 ON THE HABITS OF MOOSE.

Mr. Louis Hamel, Postmaster at Mine Centre, Ontario, to whom my

Indian sells furs. He said that, early in the winter, Billy, having

gone up the river where we counted so many moose last spring,

saw some twenty of them frozen in the ice. Evidently they had

tried to cross too early and had broken through. Perhaps no one

moose would have misjudged the strength of the ice for his own

weight; but it is rare for such large numbers to travel together.

Like men, sometimes, on a stage, they had forgotten to reckon the

combined pressure.

Such accidents, happily, are rare. There can be no doubt that -

moose are plentiful in all parts of the vast Rainy Lake District ;

for, though one may travel several days or more without seeing

the animals themselves, the signs are everywhere. If one forgets,

however, that the moose tend to congregate in certain favourite

feeding-grounds, it is easy from various record tallies to over-

estimate their total number. It is true that I saw 44 in one day

during the fly-season ; but, on the other hand, during five months’

travel the previous year, I had never seen more in a single day

than eight.

If moose were once before plentiful in this region long ago, as

some people suppose, there is no certainty that the present

conditions are permanent. Whatever swept them away before,

whether wolves, or men, or pestilence, may recur. One thing seems

sure—year by year they are pushing farther north toward the edge

of Hudson Bay. Perhaps in time by this very movement their

southern range will be deserted; but more likely they are

merely recovering old ground by force of expanding numbers.

Wolves in the Rainy Lake District are still scarce, the Indians

cannot return, and the shooting-season—perhaps unnecessarily

late—comes at a time (Nov. Ist to 15th) when the danger of

freezing lakes keeps sportsmen close to the settlements. All these

safeguards together with the ideal character of the country—its

innumerable lakes and rivers and its abundant food-supply—seem

almost to ensure the perpetuation of this noblest of American

game animals.

In the very heart of the region, too, Ontario and Minnesota

have wisely set aside contiguous tracts of more than 3500 square

miles, where the moose are protected for alltime. Thus, even with

the inevitable increase in shooting, and the probable establishment

of an earlier open season, the prospects for the future are auspicious.

It is a pleasant and noteworthy coincidence that that animal

which writers invariably describe as ‘‘ prehistoric-looking ” should

have proved among the ablest in modern times to hold its own.

3 Maia prea ae erage

Oe SE iy Pe

SEDER ee SRE I

ON SOME NEW CHINESE FLEAS. 365

PAPERS.

19. Some New Siphonaptera from China. By Karu JorDAan,

Ph.D., F.U.S., and the Hon. N. CHArRLEes ROTHSCHILD,

MATE Z.5.5 EES.

| Received and Read March 7, 1911.]

(Text-figs. 104-124.)

The following fleas were collected by Mr. M. P. Anderson in

the provinces of Shensi, Kansu, and Sze-chuen, China. ‘The

collection contains altogether 17 species, of which no fewer than

13 are new. Some of these are closely related to species described

from Turkestan or European Russia, and may possibly be only

geographical developments, others represent very distinct types

not very nearly allied to anything known from other countries.

1. ARCHHOPSYLLA SINENSIS, sp. n. (Text-figs. 104, 105.)

3 2. Agrees in both sexes very closely with A. erinacet

Bouché (1833) from Europe, differing chiefly in the following

points :—

The tooth situated in A. erinacei at the apex of the genal lobe of

the head below the antennal groove is either absent from sinensis

or small and very pale. The receptaculum seminis of the female

(text-fig. 105) is slightly smaller than in erinacei, and the modified

abdominal segments of the male exhibit some easily recognisable

characteristics as follows :—The ninth tergite, which in erinacet

has a short, broad, and curved manubrium, bears a broad and

straight manubrium with rounded apex (text-fig. 105, [X.t.). The

ventral margin of the eighth sternite is not denticulate. The large

‘movable process (1*) of the clasper is much shorter than in erimacet,

and its ventral edge, instead of being continued downwards as a

broad thin flap which is slit at the margin into filaments, bears

only a narrow membranaceous appendage. The ninth sternite

(IX. st.) is less rounded at the apex than in erirvacei and has more

bristles. As this sternite is partly concealed in our specimens

by other organs, we are not quite certain that our figure gives the

exact outline of it.

A small series of both sexes from Yu-lin-fu, Shensi, 4000 ft.,

taken off Lrinaceus miodon.

2. CERATOPHYLLUS CRISPUS, sp. n. (Text-figs. 106-108.)

3 2. The male of this species shows some very remarkable speci-

alizations not observed in any other known flea, the second segment

of the antenna bearing in that sex some very longand strongbristles

which extend far beyond the long club, and the posterior margin

of the hind coxa being incurved at some distance from the base.

Moreover, the hind tarsus of the male exhibits a development of the

bristles which is only approached to some extent in the males of

two species of the genus Vermipsylla. The female of C. crispus

366 DR. K. JORDAN AND HON. N. C. ROTHSCHILD ON

does not exhibit any trace of these specializations. The bristles

of the second antennal segment of the female are all short, and

the hind coxa and tarsus are quite normal. It agrees, however,

with the male in other characteristics—for instance, the long

rostrum and relatively short maxillary palpus, the absence of a row

i! Why Mehey

ne yy

{

]

Vilst. VIllst.

Text-fig. 104.—Clasping organs of Archeopsylla sinensis 3.

Text-fig. 105.—Abdominal segments VII & VIII and receptaculum seminis of

Archeopsylla sinensis 2.

of bristles on the inner surface of the mid and hind femora, the

presence of two rows of lateral bristles on the hind tibia, the

relatively short first mid-tarsal segment, &c.

We may mention incidentally, in connection with the drawing

SOME NEW CHINESE FLEAS. 367

of the head (text-fig. 106), that the antenna moves on a pivot

standing almost at right angles to the first segment. For that

reason the antenna cannot assume a position at right angles to

Text-fig. 106.

Head of Ceratophyllus crispus 6.

the lateral surface of the head, but, when moved from the

antennal groove, slides along the side of the occiput until the

position is reached which it has in our drawing. The hind side,

which bears the long bristles, is nearest to the eye when the

antenna is in the groove, and it is always the outer surface which

is presented.

Head.—The rostrum is a little longer in the 9 than in the

S. It reaches beyond the taodbamten | in both sexes, the apex

of the fourth segment being on a level with the base of the

trochanter and the last segment being more than twice the length

of the fourth. The maxillary palpus extends to the third

segment of the rostrum. The bristles of the head are almost the

same in number in the sexes (¢/. text-fig. 106), but are less strongly

developed in the 9 than in the J ; moreover, the thin bristles

at the anterior edge of the antennal groove are not present

in the @, and of the four bristles placed in the ¢ above the

antennal groove (1, 3) the anterior one is absent, and the large

ventral one of the row of 3 is accompanied by but one bristle,

which is small. The first antennal segment of the ¢ bears on

the inner surface a number of small bristles along the anterior

edge, and the outer surface of this segment of the 92 has

numerous small hairs. The second antennal segment of the 2

bears one row of short hairs, these hairs being very thin and only

reaching to the second segment of the club. Inthe ¢ this segment

has several very long bristles, as shown in the figure.

Thorax.—The pronotal comb consists of 18 to 20 spines. The

meso- and metanotum bear each two rows of bristles and a

number of additional short bristles which represent three more

rows on the mesonotum and two on the metanotum, being less

numerous on the latter than on the former. The mesonotum

has on the inner side eight slender spines on the two sides

together. The mesopleura bear 9 to 11 bristles, the metepister-

num 3 and the metepimerum 7 or 8 (3,3 or 4,1). There are

2 or 3 apical spines on the metanotum on the two sides

together.

“Abdomen.—The four to six anterior ter gites bear apical spines,

the numbers being (on the two sides together) i in the ¢ 2—8 or

4—9—9-_9— 0) or 2,and in the 2 2 or 3—2 to4—2—2. Theapical

edge of segments I to VII is distinctly denticulate in both sexes.

There are two rows of bristles on the tergites. The stigmata are

placed above the first bristle of the second row, and below the

first of the anterior row. ‘The 3 has one very long antepy-

gidial bristle accompanied by two short but strong ones, the upper

one being the shortest. The 2 also has free antepygidial

bristles, of which the upper one is half and the lower one two-

thirds the length of the central bristle, which is as long as the

first hind-tarsal segment. ‘The first sternite has (on both sides

together) two bristles, sternites III to VII bearing in the 3

6 bristles, and [iI to VI in the @ 6 to 8, while VII has 10 to 12

in the 2, with one or two small bristles in front of this row,

there being no additional bristles or quite exceptionally one

bristle on the other sternites.

ee ae eee ee ek ee

SOME NEW CHINESE FLEAS. 369

Legs.—The hind margin of the hind coxa of the ¢ is incurved

from one-fourth of the margin to its centre. The mid- and

hind femora bear on the inside one lateral bristle, placed at the

basal third, and on both sides one subapical ventral bristle. The

hind tibia has a double row of 13 to 16 bristles on the outer

surface and a single row of 5 to 7 on the inside. The outer

dorsal bristles, 18 in number inclusive of the apical one, are stout,

most of them being less pointed than usually and nearly all being

of more or less the same length. The longer bristles have like-

wise blunt tips. The longest apical one hardly reaches to the sub-

apical notch of the first hind-tarsal segment. The first segment

of the mid-tarsus is distinctly shorter than the second. The

fourth segment of the fore- and mid-tarsi is only as long as it is

broad, that segment of the hind tarsus being one-tenth longer than

broad in the Q and about one-fifth in the ¢. The fifth segment

bears in all the tarsi five pairs of lateral bristles. The hind tarsus of

the ¢ is remarkable for the peculiar development of the bristles.

The bristles on the upper surface and at the hind edge of these

segments are thin and those placed in the notches of the hind edge

and at the apex are long, the longest bristles, moreover, being wavy.

There are eight bristles each dorsally at the apex of the second

and third segments. These bristles radiate in fan-shape, the one

placed at the anterior apical corner deviating but little from the

general direction of the tarsus and being the shortest of the row,

the others gradually increasing in length as one proceeds from

the anterior to the posterior side of the segment, the longest

about equalling segments 3, 4,and 5together. The bristles on the

tarsi of the 9 are all short, the apical ones of the second hind-

tarsal segment not reaching even to the apex of the third segment.

The proportional lengths of the segments are as follows :—

Midi tarsus: Glee ZON LIE (elo OF 1, 20 etl 7. NO!

Modified segments. The eighth tergite bears about 20

bristles in the upper half of the apical lobe and none in the lower

half. The eighth sternite (text-fig. 107, VIII. st.) is narrow and

curved, and bears ventrally two bristles. Its distal portion is

membranaceous and divided into a large fringed flap and several

long filaments as indicated in the figure. The clasper (text-fig.

107, Cl) is small and is produced into a vertical, slightly club-shaped

process (P), which bears two small hairs. The movable process

(F), on the other hand, is very large and presents two remarkable

structures. The upper edge, is, as far as we can make out,

widened into a membranous appendage which projects distally

and does not bear any hairs. The other peculiarity is the

presence of a row of short spine-like bristles on the outer surface

of the “finger.” ‘The ninth sternite (text-fig. 107, IX. st.) hasa

broad, curved vertical arm and a long and relatively narrow

horizontal arm, both being of a characteristic shape. The distal

portion of the horizontal arm narrows almost to a point.—

370 DR. K. JORDAN AND HON. N. C. ROTHSCHILD ON ~

@. The apical margin of the seventh sternite is slanting and more

or less undulating, the two sides not being exactly alike. The

ventral angle of this sternite projects more or less strongly and is

sometimes almost pointed (text-fig. 108, VII. st.). The eighth

tergite (VIII. t.) bears on each side only 4 to 7 small bristles above

the stigma, and below it one long one accompanied by one to four

small ones. There are 12 bristles on the lower half of the

segment as shown in the drawing (text-fig. 108). The apical

margin is angulate below the centre and the upper angle more or

Text-fig. 107.

Clasping organs of Ceratophyllus crispus 3.

less rounded off. The ninth sternite—more or less membranaceous

and lying inside the eight tergite—bears some small hairs. The

stylet is almost cylindrical and bears one bristle beside the apical

one. ‘The anal sternite has 4 very long bristles at and near

the apex and proximally to them, on the underside, about 6 heavy

spine-like bristles, besides a number of short lateral and ventral

bristles, of which some are thin and others stout. The recepta-

culum ces (text-fig. 108) is slender, the head being shorter than

the tail.

SOME NEW CHINESE FLEAS. 371

Length (mounted specimens) 2°4 to 2°8 mm.

Two males and four females from Omi-shan, Sze-chuen, 6000 ft.,

off Scturotamias davidianus consobrinus M.-Edw., and from 23

miles 8.K. of Ta-tsien-lu, 7500 ft., off the same host.

Text-

fig. 108.

Text-

fig. 109.

Text-fig. 108.—Abdominal segments VII & VIII, stylet and veceptaculum seminis

of Ceratophyllus crispus 2.

Text-fig. 109.—Abdominal segments VII & VIII and stylet of Ceratophyllus

dolabris 2.

3. CERATOPHYLLUS DOLABRIS, sp. n. (Text-figs. 109, 110.)

3 2. Nearly allied to C. lagomys Wagn. (1897), of which only

the ¢ is known to us. The new species is distinguished from

that sex of lagomys by the shorter bristles of the eighth abdominal

sternite, the differently shaped non-movable process of the

clasper, the much narrower distal portion of the ninth sternite, the

broader lobes of the anal sternite, and some other details men-

tioned below. The most characteristic feature in both lagomys

and the new species is the development of the tenth abdominal

sternite of the ¢. This sternite consists in fleas normally of a

aly DR. K. JORDAN AND HON. N. C. ROTNSCHILD ON

single sclerite. Im the males of the species under discussion,

however, it is completely separated in the mesial line into a right

and left lobe, which are very long and slender, projecting far

beyond the tenth tergite, and bearing numerous bristles on the

upper and lateral surfaces but none on the underside.

Head.—The frontal tubercle is prominent. There is a row of

3 long bristles in front of the eye. The occiput bears one long

bristle above the antennal groove, besides the subapical row of

bristles. The bristles on the second segment of the antenna of

the 9 are long, there being at least five which reach beyond the

club. The rostrum extends to the base of the femur and the

maxillary palpi reach the trochanter.

Thorax.—The comb of the prothorax consists of 17 or 18 spines,

there being usually an additional small spine on each side. The

meso- and metanotum have on the two sides together a posterior

row of 12 bristles, and before it a row of about 6, the mesonotum

bearing in addition a row of hairs at the base and several short

bristles on the back. The metanotum has four apical spines on

the two sides together. There are from 4 to 6 bristles on the

metepimerum (1 or 2, 2 or 3, 1), usually 4 or 5.

Abdomen.—The tergites bear each two rows of bristles, the

first tergite having 2 or 3 additional bristles in front of the rows.

On none of the tergites does the anterior row extend down to the

most ventral bristle of the posterior row, the row stopping short

at the third or fourth bristle of the posterior row on the central

segments in the ¢, and at the second or third in the Q.

Tergites I to IV or V bear apical spines like the metanotum,

the numbers being on the two sides together, 4—4 to 6—4—2 to

40 to 2. The ¢ bears one long antepygidial bristle accom-

panied by two minute hairs, while the Q has 3 bristles (all

broken on both sides in our only 2). The bristles on the

sternites number on the two sides together in the ¢ 2, 6, 7, 8, 8,

8, and in the Q 5, 9, 11, 11, 11, 15, there being 16 additional

bristles on the seventh sternite of the 2, which are placed

irregularly in front of the row. The stigmata in the 2 are placed

a little above the first bristle of the posterior row of the tergites

or on level with it, in the ¢ below it on the posterior segment,

Legs.—The hind femur bears on the outer side one subapical

ventral bristle and in the anterior half one or three lateral ones.

On the inner surface there is a row of 9 to 12 bristles. The hind

tibia has outside a row of 8 or 9 lateral bristles and on the inside

a row of 4to 7. The longest apical bristle of the second hind-

tarsal segment reaches beyond the apex of the third. The fourth

hind-tarsal segment is nearly twice as long as it is broad near the

apex.

P Modified segments.— ¢. The eighth tergite has 3 bristles below

the stigma, a row of 6 to 8 at the upper edge of the apical lobe,

with 1 or 2 bristles close to the row, and an oblique and more or

less irregular row of 5 or 6 near the ventral edge. The eighth

sternite (text-fig. 110, VIIL. st.) is similar to that of C. lagomys.

SOME NEW CHINESE FLEAS, 373

It bears, as in that species, three bristles at the apex on each

side. While the two longest of these bristles are, however, half

the length of the sternite in Jagomys, they measure only about

one-third the length of the sternite in the new species. The

clasper is much the same as in C. lagomys, but the process (P)

(text-fig. 110) is much more incurved on its distal side and hence

the lower corner of the widened apical portion more pointed.

The upper edge of the triangular movable process (F'} (text-fig. 110)

is less rounded than in lagomys, the two spines are pointed, and

the long bristle of the distal edge is placed farther away from

Text-fig. 110.

Weer) iis

‘ Va Ne

Clasping organs of Ceratophyllus dolabris 6.

them than in lagemys. The ninth sternite (text-fig. 110, LX. st.)

is characterised, as in C. lagomzs, by the central ventral lobe of

the horizontal arm bearing a number of rather strong, short

bristles at the apex. The thin, pale, distal lobe is more rounded

than in lagomys and much broader.— 9. The seventh sternite

(text-fig. 109, VII. st.) is truncate and slightly incurved twice.

The eighth tergite (VIII. t.) bears 5 to 6 bristles below the stigma

in two rows and has the upper angle of the apical lobe strongly

rounded, the centre of the apical margin being somewhat tri-

angularly produced as shown in the figure. The bristles of the

Proc. Zoou. Soc.—1911, No. XXV. 25

374 DR. K. JORDAN AND HON. N, C. ROTHSCHILD ON

stylet are long, the one placed near the ventral edge (side-view)

being exceptionally long.

Length (mounted specimens) 2°5 to 3°2 mm.

Three males and one female from Old Tau-chow, Kansu, 9000 ft.,

off Marmota robusta M.-Kdw.

4. CERATOPHYLLUS CRASSUS, sp. n. (Text-fig. 111.)

The only specimens (two females) which we have were in the

same tube as the specimens of C. dolabris, being found on the

same host, presumably on the same individual of the host. They

resemble the female of C. dolabri is, but are much more hairy and of

stronger build.

Heatdl.— The frons bears a row of three bristles before the eye,

the upper two standing rather close together and there being a

fourth bristle in front of the ventral one of therow. The occiput

bears a moderately long bristle above the antennal groove, about

10 small hairs along the groove, one of them being longer and

thicker, and the usual subapical row. The two lower bristles of

this row are large and there is a wide interspace between them

and the next bristle of the row. The bristles of the second

antennal segment are long, at least five of them reaching the

apex of the club. The frontal tubercle is prominent. The

maxillary palpus extends to near the apex of the fore coxa,

while the rostrum reaches far beyond the trochanter, the apex

of the fourth segment being on a level with the base of the

trochanter.

Thorax.—The pronotal comb consists of 20 to 23 spines. The

meso- and metanotum bear each two rows of bristles. The

bristle-like spines on the inner surface of the mesonotum are

very numerous (14 to 16 on the two sides together).

Abdomen.— The tergites bear three rows of bristles, the anterior

row being more or less irregular, the second reaching down tothe

stigma and the third row having 2 or 3 bristles placed below the

stigma. The tergites | to IV or V bear apical spines like the

metanotum, the numbers on the two sides together being 4—5

to 7—4 to 6—2 to 5—0 or 1. The edges of the tergites are

irregularly excised dorsally, but do not show any distinct minute

serration as is usually the case in the allied forms. There are

three antepygidial bristles (broken in our two specimens). All

the sternites have additional bristles in front of the usual

posterior row, the numbers of bristles being as follows on the two

sides together (the first number referring to the additional

bristles of each SeRMEBD) : 5 or 6, 6 te: Bas 19 to. 21, 16 or 17—22

to 24, 15 to 17—22 to 26, 14 to es 22, 15—18 to 22 poe

Legs, —The wena femur bears on the outside a row of 5 or 6

bristles and on the inside a row of 11 or 12. The hind tibia has

11 lateral bristles on the outside and 8 on the inside. Two of

the bristles of the second hind-tarsal segment reach beyond the

apex of the third. The fourth segment in all tarsi is only one-

‘s

SOME NEW CHINESE FLEAS. 375

third longer than it is broad near the apex. The measurements

of the tarsi are :—

Mid tarsus: 28, 24, 20, 13, 30.

Hind tarsus: 58, 36, 26, 17, 33.

Text-fig. 111.

Ash

Abdominal segment VIII and receptaculum seminis of Ceratophyllus

crassus 2.

Modified segments.— 9. The seventh sternite is so much

torn in both our specimens that it is not safe to express an

opinion about its outline. The eighth tergite (text-fig. 111) bears

two long bristles below the stigma and its apex is obliquely

truncate and slightly incurved. The bristles on the lower half

of the eighth tergite are numerous, and there are 3 or more short

strong bristles on the inner surface, The stylet resembles that

of C. dolabris, sp. n., but the apical bristle is appreciably shorter

and thinner. The receptacle has a rounded head, which is shorter

than the (artificially ?) distorted tail,

Length (mounted specimens) 3°8 to 4 mm.

Two females from Old Tau-chow, Kansu, 9000 ft., off Warmota

robusta M.-Kidw.

5. CERATOPHYLLUS MANDARINUS, sp. n. (Text-figs. 112, 113.)

3 2. A pale species with long rostrum, long tarsal bristles,

a very long and narrow movable process in the male genitalia,

and a peculiarly long-tailed receptaculum seminis with a small

head in the female.

Head.—The frons is very feebly rounded in both sexes, the

25%

376 DR. K, JORDAN AND HON. N. C. ROTHSCHILD ON

frontal tubercle being placed at one-third or one-fourth the

distance from the oral corner to the antennal groove. ‘There are

two long bristles in front of the eye, one of them standing near

the eye and being accompanied by a much shorter bristle, and

the others being placed at the oral margin. Further frontad

there are two more bristles in the ¢, but not in the 9, one of

them at the oral margin behind the maxillary palpus and the

other near the antennal groove. On the occiput there is one

bristle above the antennal groove, and along the groove about

half-a-dozen small hairs in both sexes. The apical row of bristles

contains 1 long and 3 or 4 small ones, the interspace between the

long (ventral) bristle and the next one being large. The third

segment of the maxillary palpus is comparatively long, being only

one-eighth shorter than the second segment. The rostrum

reaches well beyond the trochanter, the proportional lengths of the

seements being, 16, 15, 15, 27, and 40.

Thorax.—The pronotum has a comb of 18 spines (in one of the

females 23) and a row of 12 bristles on the two sides together.

On the mesonotum the postmedian row consists also of 12 bristles,

while the row before it contains from 8 to 10. Near the base of

the mesonotum there are only 14 small hairs in the g and 24 in

the 9, while the inner surface bears near the apex from 9 to

12 bristle-like spines. The mesopleura bear five long bristles and

anteriorly a few small hairs in addition. ‘The numbers of bristles

on the metanotum are 12 in the postmedian row in both sexes, 3 to

5 in the anteriorrowinthe ¢ and9tollinthe 2. The metepi-

sternum bears 2 or 3 bristles and the epimerum 5 (2, 2, 1).

There is on each side one apical spine on the metanotum, as on

the first to third or fourth abdominal tergites.

Abdomen.—The tergites I to VII bear in the 2 each two rows

of bristles, the postmedian row containing 13 or 14 bristles on the

central segments; in the ¢ the anterior row is reduced to 4 to 6

bristles on the first to fourth segments and to one or two on

segments V and VI. The seventh tergite bears on each side in

the ¢ one long antepygidial bristle accompanied by two minute

hairs, and in the @ two long bristles, of which the lower one is a

very little shorter than the other. The numbers of bristles on

the sternites are in the ¢ 2—4 to 6—6—6—4, and in the 9?

2—12 to 14-8—7 to 10—8—13 or 14, there being 3 or 4 small

bristles in front of the row of the seventh segment on the two

sides together.

Legs.—The hind femur has on the outside two Jateral bristles

and one subapical one and on the inside a row of 7 or 8 bristles.

On the hind tibia there is one lateral row on the outside,

containing in the ¢ 7 bristles and in the 2 5 to 8, the inner

surface bearing a row of 4 to 7 in the ¢ and of 6 to 8 in

the 2. ‘The first mid-tarsal segment is the same in length as the

second. The longest bristle of the first hind-tarsal segment

reaches at least to the apex of the second, and this latter segment

bears two apical bristles which extend considerably beyond the

SOME NEW CHINESE FLEAS. 377

apex of the fourth. The proximal pair of bristles of the fifth

segment is lateral like the other pairs. | The proportional lengths

ot the tarsal segments are :—

Horetarcsus LO; 12°108 99)" OF Ty ts 12, 10, 24.

Widttamsass) 18, 18s L459," 2D. 0 24 Ooms Or aey

Hind tarsus: ¢. 49, 32, 20,12, 24; 9. 60, 37, 23, 14, 30:

Text-fig. 112.

Clasping organs of Ceratophyllus mandarinus @.

Modified segments.— ¢. The eighth tergite bears at the upper

edge beyond the stigma a row of four long The istles, from the last

bristle towards the "ides of the segment there are two bristles,

another stands below the stigma and two are situated near the

ventral edge. . The eighth sternite (text-fig. 112, VIII. st.) issmall,

being distally suddenly narrowed to a point in lateral aspect and

bearing no bristles. The clasper (text-fig. 112, Cl) is broad, its

upper angle rounded and not produced into a process. The

manubrium is at first strongly narrowing from the clasper inward

and then remains almost of even width: its apex is slightly

eurved downwards and has the upper angle more strongly

rounded off than the lower angle. ‘The movable process (F) is

very long and of practically ene same width throughout. It is

but slightly curved and its tip rounded off. This finger bears a

short bristle at the tip, a long one below the tip at the distal or

ventral margin, and a third short bristle further down, then follow

another short one, a moderately long one and another a little

shorter than the preceding one. This last bristle is. situated a

short distance heyond the centre of the margin of the finger, there

378 DR. K. JORDAN AND HON. N. C. ROTHSCHILD ON

being in one of the specimens another bristle lower down. The

vertical arm of the ninth sternite (IX. st.) is very slender, especi-

ally the upper portion, which les at the manubrium of the

elasper. The ventral arm is divided by a ventral sinus into a

longer proximal portion and a shorter and broader distal portion,

the latter being about as long as it is broad. The proximal

portion bears at the ventral margin 5 or 6 bristles, the two distal

ones of which stand close together one on each side, and are fairly

stout in comparison with their length. 9. The seventh sternite

Text-fig. 113.

MIL.

Villst

“NILSt

Abdominal segments VII & VIII and receptaculum seminis of

Ceratophyllus mandarinus 2 .

(text-tig. 113, VII, st.) narrows in side-view ; its apex is truncate,

somewhat obliquely, and slightly emarginate, with the upper

apical angle qnite distinct, The eighth tergite bears 8 to 10

small bristles above the stigma, 2 or 3 long ones beneath the

stigma besides some small ones, and there are about 12 long

bristles and 10 to 12 shorter ones on the lower half of the segment.

The stylet is twice and one-half as long as broad. ‘The bristles

on the ninth and tenth tergites (which segments are separated by

a distinct suture) are numerous. The receptaculum seminis is

distinguished by the head being nearly three times as wide as the

tail and only half its length, It has been disturbed in its position

by the mounting in both specimens which we have, and the out-

line we give does not exactly represent a true lateral aspect of the

organ.

“Phree males and two females from Yu- lin- fu, Shensi, 4000 ft.,

taken off Citellus mongolicus.

RENEE gg Dig 24

SOME NEW CHINESE FLEAS, 379

6. CERATOPHYLLUS MONGOLICUS, sp, n. (Text-fig. 114.)

3 2. Asingle pair in the collection agrees well with C. tes-

quorum Wagn, (1893), except in the details mentioned below.

The new species is smaller and paler than C. tesqworwm and bears,

on the whole, fewer bristles. The rostrum reaches a little beyond

the trochanter in both sexes and is therefore longer than in the

species mentioned. The abdominal sternites III to VI have each

a row of 6 bristles on the two sides together, the 2 bearing one or

two additional bristles on each side of the sixth segment. The

hind femur has on the outside one ( 3 ) or two (Q ) lateral bristles in

the basal half and a subventral bristle before the apex, and on

the inner side a row of 8 or 9 bristles. The hind tibia, which has

a double row of about 11 bristles on the outer surface in C. tes-

quorum, has only 8 lateral bristles in C. mongolicus inclusive of

the apical one. The first mid-tarsal segment is distinctly shorter

than the second,

Text-fig. 114.

Clasping organs of Ceratophyllus mongolicus 3.

The differences in the male genitalia are slighter than they

generally aye in distinct species, The non-movable process of

the clasper (text-fig. 114, P) is a little slenderer than in (C. tes-

quorum. The finger (F) is triangular as in that species, and bears

the same bristles, but it is less widened at the apex, the upper

(=apical) margin being one-fourth shorter than in tesguorwm,

the proportions of length and width of the finger being 7:4 in

mongolicus and7:5in tesquorum. The eighth sternite of tesquorwm

is provided with 5 bristles and a thin hair, the bristles being

380 DR. K. JORDAN AND HON, N. C. ROTHSCHILD ON

short but fairly thick; in the new species there are some weak

hairs instead (text-fig. 114, VIII. st.).

@. The seventh sternite of the 2 bears on each side a row of

12 or 13 bristles and about 6 small bristles in front of the row.

The apex ot this sternite (in lateral aspect) is truncate, the

margin being very slightly excurved centrally. The eighth

tergite has a cluster of 5 or 6 bristles below the stigma and about

15 bristles at, and near, the ventral and apical margins. As in

the 2 of C. tesquorwm, there are two long antepygidial bristles on

each side, not accompanied by a third stout bristle, and the

bristles of the anal sternite stand far removed from the base in

a dense cluster occupying the apical two-fifths of the sternite.

One pair from Yu-lin-fu, Shensi, 4000 ft., taken off Citellus

mongolicus.

7. CHRATOPHYLLUS FAMULUS, sp. n. (Text-fig. 115.)

@. Closely resembling the preceding species, but the bristles

on the body and legs are more numerous.

Head.—Frons moderately curved, slanting. The tubercle small,

placed a little nearer to the central sensory organ (pale dot) than

to the oral corner, In front of the well-developed eye there is a

row of three bristles, the upper one being placed a little lower

than the centre of the eye and the second, being nearer to the

upper than to the lower bristle, | Between the upper bristle and

the antennal groove there are several minute hairs. The occiput

has above the antennal groove one long bristle and about 14

minute hairs. The subapical row of bristles of the occiput

contains on each side five or six, of which the lowest is the

largest, being separated from the next bristle by a wide interspace.

The bristles of the second antennal segment are long, ten of them

reaching the apex of the club ory beyond. The rostrum extends

beyond the base of the fore femur, the apex of the fourth segment

being on a level with the base of the trochanter. The maxillary

palpus reaches to the base of the fourth segment of the rostrum.

Thorax.—The pronotal comb ccnsists of 20 spines. The meso-

and metanotum have each two rows of bristles, the second row

containing 11 or 12 and the anterior one 15, on the two sides

together. The mesonotum has about six and the metanotum two

or three additional bristles on the back. The mesopleura have

about five long bristles and anteriorly several thin hairs. The

metepimerum bears six bristles (2, 3, 1). There are two short

apical spines on each side of the metanotum.

Abdomen.— The tergites have all two rows of bristles, with one

or two additional bristles on the back; tergites I to IV hearing,

moreover, some apical spines. The stigmata are placed above the

first bristle of the posterior row, but far in front of it. The first

sternite bears a pair of ventral bristles, and no lateral bristles or

only one. The sternites of segments III to VI have on the two

sides together a postmedian row of 8 bristles and a few additional

small bristles in front of therow, The seventh segment, however,

SOME NEW CHINESE FLEAS. 381

has ten long bristles in a row and no less than about 30 additional

bristles, on the two sides together. “There are two long antepy-

gidial bristles on a double cone.

Legs.—The fore femur has on the outer surface about 11 lateral

bristles arranged in two irregular rows and on the inside a row of

five lateral bristles. ‘The mid and hind femora have outside one

or two lateral bristles, and inside a row of 9 or 10 and 12 or 13

respectively. ‘The dorsal and apical bristles of the tibiz and the

apical ones of the tarsal segments are stout. The hind tibia has 10

lateral bristles on the outside and 5 to 7 on the inside. The

longest apical bristle of the mid tibia almost reaches to the apex

of the second tarsal segment. ‘The second hind-tarsal segment has

two apical bristles which extend to the apex of the third segment

or beyond. The fifth tarsal segment has five lateral pairs of

bristles. The measurements of the tarsal segments are as

follows :—

Mid tarsus: 20,21, 14, 9, 22,

Hind tarsus: 53, 34, 21, 12, 24.

Text-fig. 115.

Abdominal segments VII & VIII and receptaculum seminis of

Ceratophyllus fumulus § .

Moiified segments.— 2. The apical margin of the seventh

sternite (text-fig. 115, VII. st.)is slanting and slightly undulating.

The eighth tergite bears two rows of bristles beneath the stigma

and about 16 bristles on the lower half. The apical margin of

this tergite is rounded-truncate, with the upper angle distinctly

produced. The ninth sternite (not drawn) bears several small

bristles. The stylet, which has the shape of a champagne bottle,

is twice as long as it is broad. The head of the receptaculum

seminis is about as long as the tail.

Length (of mounted specimens) 2°4 to 2°6 mm.

Two females ; locality and host as in species 3 and.4.

382 DR. K. JORDAN AND HON. N. GC. ROTHSCHILD ON

8. CmrrATOPHYLLUS EUTELES, sp. n. (‘Text-fig. 116.)

@. This is one of the species in which the first pair of bristles

of the fifth segment of all the tarsi is shifted on to the ventral

surface, being placed almost in between the second pair. The

species may be recognised by the absence of bristles from the first

abdominal sternite, Sine shortness of the bristles on the tarsi, the

paucity of bristles on the femora, &e.

Head.—The frontal tubercle is placed nearer to the central

sensory organ (pale dot) than to the oral angle. The eye is well —

developed and pigmented. There is a row of three long bristles

before the eye, the uppermost bristle being a little lower than the

centre of the eye. In front of this bristle there are one or two

small bristles and usually a third further upwards nearer the

antennal groove. ‘The occiput bears a large bristle above the centre

of the antennal groove, a small bristle further frontad and another

small one familie dorsad, The subapical row of bristles of the

occiput consists on each side of 6 bristles, of which the ventral one

is large. The interspace between this bristle and the next is twice

the size of the interspace between the second and third bristles of

that row. Five of the bristles of the second antennal segment are

prolonged, two of them nearly reaching to the apex of the club.

The rostrum extends to the apex of the coxa, its fifth segment

being more than twice the length of the fourth. The maxillary

palpus reaches to the apex of the fourth segment of the rostrum,

the proportional lengths of its segments being 17, 15, 12, 19.

Thorax.—The pronotal comb consists of 18 spines. The meso-

and metanotum bear each two rows of bristles and a few additional

dorsal bristles representing a third row. The mesopleura have 8

bristles and some additional short stout hairs. The metepimerum

bears 6 bristles (2, 3, 1). The metanotum has one apical spine on

each side.

Abdomen.—The tergites bear each two rows of bristles as

follows (the first number being that of the anterior row): 8 or 9,

9—8 to 10, 12—8 to 10, 12—7 to 9, 12 or 13—5 or 9, 12 or 13—

5 or 8, 11 or 12—3 or 4,9 or10. The first sternite has no bristles

at all. The sternites of segments III to VI have on each side

three bristles, and that of segment VII bears four bristles

with an additional bristle in front of the row. There are three

antepygidial bristles, the middle one being twice the length of the

upper and one-fifth longer than the lower.

Legs.—The fore femur has two bristles on the outer surface, and

one on the inner, apart from a ventral subapical bristle. The mid

and hind femora have only the subapical ventral bristle on the

outside, and on the inside a small lateral bristle and a small ven-

tral subapical one. The hind tibia has a row of 8 lateral bristles

on the outside and two or three bristles on the inner. The first

and second hind-tarsal segments are long and slender and the

bristles of the tarsi short. The longest apical bristle of the second

hind-tarsa] segment is one-third shorter than the third segment.

The fifth segment is rather short in all the tarsi and bears

can

SOME NEW CHINESE FLEAS, 383

ventrally at the apex two short and fairly slender bristles, which

are placed far apart. The first lateral pair of this segment stands

nearly in between the second pair. The measurements of the

tarsi are :—

Mid tarsus: 26, 23,15, 9, 20.

Hind tarsus: 54, 34, 23, 13, 21.

Modified segments.— 2. The seventh sternite (text-fig. 116)

rather strongly narrows apically, being truncate-sinuate in two

specimens as shown in the figure, while in a third the apical margin

Text-fig. 116—Abdominal segemnts VII & VIII and receptaculum seminis of

Ceratophyllus euteles 2.

Text-fig. 117.—Abdominal segments VII & VIII of Ceratophyllus pheopis.

is more slanting and hence the ventral angle (in side view) more

pointed. The eighth tergite (VIII. t.) bears one long and two short

bristles below the stigma and 7 to 10 bristles in the ventral half,

three of which are placed at the edge above the ventral angle.

384 DR. K. JORDAN AND HON. N. C. ROTHSCHILD ON

The apical margin of this tergite is gently incurved, its upper

angle being rounded, and the lower one projecting and being

almost pointed. The ninth sternite (1X. st.) bears one short but

rather stout bristle at itslower corner. The receptaculum seminis

has an elongate head, which is longer than the tail.

Length (mounted specimens) 2°4 to 2°7 mm.

Three females from 23 miles S.E. of Ta-tsien-lu, 7500 ft., off

Sciurotamias davidianus consobrinus M.-Kdw.; Omi-shan, Sze-

chuen, 9500 ft., off the same host and off Tamiops swinhoei

M.-Edw.

9, CERATOPHYLLUS PHMOPIS, sp. n. (Text-fig. 117.)

Q. This species, of which we have only one female, is easily

recognised hy the very feebly pigmented eye, the very short bristles

of the antenna, and the deeply sinuate seventh abdominal sternite.

Head.—The frons is very slanting. The frontal tubercle stands

about halfway between the oral corner and the central sensory

organ (pale spot), the distance from the tubercle to the oral angle

being about equal to the diameter of the eye. There are three

long bristles in front of the eye, the uppermost being placed near

the antennal groove, the interspace between this bristle and the

second being nearly twice as large as the interspace between the

second and third bristles. In front of the lower bristle there is

one small bristle, several minute hairs being placed in between the

two other bristles. The occiput bears one bristle above the

antennal groove, and a subapical row of 5 (on each side), the most

ventral of them being longand strong. The bristles of the second

segment of the antenna are quite minute. The rostrum reaches

to the apex of the coxa, the last segment being half as long

again as the fourth. The eye is feebly pigmented, excepting

the anterior and posterior edges. It appears deeply excised at a

certain focus.

Thorax.—The comb of the pronotum consists of 18 spines, the

most ventral one of one side being very small. The meso- and

metanota bear each two rows of bristles. The mesopleura have 5

or 6 bristles and the metepimerum 8 (4, 3,1). The metanotum

has also 2 apical spines on each side.

Abdomen.— The tergites bear two rows of bristles, the anterior

row containing but a small number of them, the numbers in the

two rows being 7 and 14 on the fourth. tergite on the two sides

together. The stigmata are placed dorsally to and in front of the

ventral bristle of the second row. There are three antepygidial

bristles, the lower one being but little shorter than the central

bristle, while the upper one is about one-third the length of the

latter. The basal sternite has on each side a patch of four lateral

bristles and ventrally one bristle. The numbers of bristles of the

sternites of segments III to VII are on the two sides together

(the first number giving the bristles placed in front of the row) 4,

11—3, 11—2, 8—0, 10. Tergites I and II bear an apical spine

on each side.

SOME NEW CHINESE FLEAS. 385

Legs.—The hind femur bears on the outside one subapical ventral

bristle and one lateral subbasal one, and on the inside a rew of 5 or

6 bristles. The hind tibia has a single lateral row of 7 or 8 bristles

on the outside and a row of 3 or 4 on the inside. ‘The tarsi are

slender. The second hind-tarsal segment bears an apical bristle

which reaches beyond the apex of the fourth segment and another

which extends beyond the apex of the third. The fifth segment

ears in all the tarsi five pairs of lateral bristles. The measure-

ments of the tarsi are :—

Mid tarsus: 29, 25, 15, 11, 2

Hind tarsus : 58, 38, 23, 13, 2

Modified segments.— ? . The seventh sternite (text-tig.117, VII.

st.) bears a deep sinus and is strengthened proximally to the sinus

by a curved band-like incrassation. The eighth tergite has three

bristles below the stigma, arranged in a triangle. The bristles on

the lower half of the segment are fewin number. The apical

margin is rounded and, above the marginal bristles, slightly

incurved. The ninth sternite (IX. st.) bears a few minute hairs.

The stylet is short and bottle-shaped, being twice as long as it is

broad near the base. Its apical bristle is a little smaller than the

most ventral one of the seventh tergite. The receptacle stands on

end in our only specimen, and for that reason its shape cannot well

be made out. The head appears to be round, and much shorter

than the tail.

Length (of mounted and extended specimen) 3°2 mm,

One female from 23 miles 8.E. of Ta-tsien-lu, 7500 ft., off

Sciurotamias davidianus consobrinus M.-Kdw.

10. AMPHIPSYLLA CASIS, sp.n. (Text-figs. 118, 119.)

3 2. The nearest ally of this species is 4. dea Dampf (1910)

from Turkestan. The detailed description which A. Dampf gave

applies almost verbally to the present flea, except for the points

mentioned below, and the beautiful flour es published by that

author of the head of the ¢ of dea anal the last segments of the

Q agree also equally well with the new species. As in some other

instances mentioned in this paper, we suspect that we are dealing

with a Chinese (or Hast-Asiatic) representative of dea.

Both sexes have, on the whole, less bristles than 4A. dea. The

sternum of the metathorax bears only one long bristle, which is

occasionally accompanied by a minute hair. The stigmata of the

abdomen are situated between the second and third bristle of the

posterior row, not above the third, on the seventh segment some-

times below the second bristle.

The epimerum of the metathorax bears 7 bristles only (2, 3, 2).

The anterior row of bristles on the first abdominal tergite con-

tains on the two sides together about 8 bristles, and the posterior’

row 10, the numbers of bristles in the posterior rows on the other

tergites being (also on the two sides together) 18, 17, 16, 16,

eG:

386 DR. K. JORDAN AND HON. N. C. ROTHSCHILD ON

The bristles on the sternites II to VII number on the two

sides together 2, 8, 7, 7, 8,9, there being no short bristles in front

of the row. The eighth tergite is similar to the figure given by

Dampf of that segment of 4. daa, but the bristles near the upper

edge are more numerous and those on the side less numerous.

The eighth sternite (text-fig. 118) is more densely hairy than in

A. dea, the hairs at the inner side of the margin being very

numerous, thin and more or less wavy, while those on the outer

side are longer and thicker, the longest hairs being longer than

the eighth sternite. The clasper (Cl) is less broad than in

Dampf’s figure and its upper margin more distinctly incurved,

Text-

fig. 118.

Toxt- Seay Milt

fig. 119. Sn gt hak

ils Villst,

Text-fig. 118.—Clasping organs of Amphipsylla casis 6.

Text-fig. 119.—Abdominal segments VII & VIII and receptaculum seminis of

Amphipsylla casis ?.

the portion beyond the insertion of the movable process being

consequently slenderer thanin A.dewa. The finger itself (I) bears

the same armature as in A, dwa, but is altogether slenderer, and

at the apex distinctly narrowed, not widened as in 4.dea. ‘The

ninth sternite (IX. st.) is not at all the same asin A. daa. The

apical third of the horizontal arm is very narrow, its top 1s more

strongly chitinized, and bears a short, proximally thick, terminal

bristle on each side; the distribution of the fine hairs may be

gathered from the figure.

The bristles of the abdomen and legs are more numerous than

SOME NEW CHINESE FLEAS. 387

in the ¢, their number coming within the limits of variation of

dea as given by Dampf.

The receptaculum seminis differs from Dampf’s figure of that

organ of A. dea in having a distinctly shorter tail.

One male and two females from Y u-lin-fu, Shensi, 4000 ft., taken

off Myospalax fontaniert.

11. NeopsyLua compar, sp. n. (Text-fig. 120.)

2. Verysimilar to VW. bidentatiformis Wagn. (1893), from which

it differs chiefly in the following points:—The pronotum bears a

row of 8 small bristles in front of the postmedial row of bristles

on the two sides together. The seventh abdominal sternite is a

little more deeply sinuate (lateral aspect) and its bristles are rather

stronger, The hind coxa, which bears in bidentatiformis a large

number of slender hairs on the inner side along the anterior edge,

has a patch of short spines besides many hairs. The dorsal

bristles of the hind tibia are thinner in compar than in bidentati-

formis, and the hind tibia bears on the outer surface two rows of

only 9 instead of 11 bristles, the rows, moreover, being separated

from the ventral bristles by a wide interspace without bristles,

Text-fig. 120.

VTL,

“AVMs Villst

Abdominal segments VII & VIII and receptaculum seminis of

Neopsylla compar &.

while in bidentatiformis the interspace bears several additional

bristles. The bristles at the hmder side of the first segment of

the mid and hind tarsi are thicker than the proximal bristles of

the anterior edge of the segment in Wagner’s species, there being

three such bristles in the mid tarsus and five in the hind dereme,

apart from the apical bristle. In the new species the bristles of

these tarsal segments are practically the same on the two sides.

Moreover, there are 10 or 1] bristles on the outer surface of the

first hind-tarsal segment of lidentatiformis and only 8 or 9 in

compar. Thesecond mid-tarsal segment is a little shorter in compar,

the measurements of the first and second segments being in

compar 28 and 19, and in bidentatiformis 28 and 23. The eighth

tergite has in compar on the outer surface a marginal row of 6 or

388 DR. K. JORDAN AND HON. N. GC. ROTHSCHILD ON

7 bristles, and near these bristles about 8 more, of which 2 are

long (text-fig. 120). In bidentatiformis the marginal row contains

9 bristles and there are about 15 lateral ones, of which at least

4are long. In both species is the head of the receptaculum seminis

half as long again as broad, and the tail half as long again as the

head. The two insects do not present any appreciable difference

in this organ.

One female from Yu-lin-fu, Shensi, 4000 ft., taken off Dignus

sowerbyi.

12. NEOPSYLLA ALIENA, sp.n. (Text-figs. 121, 122.)

3 2. Differs from true Weopsylla in the hind coxa bearing a

patch of short spines on the inner surface, and in the fifth segment

bearing in all the tarsi five lateral bristles. Both characters are

of great interest. The development of bristles into short spines

on the hind coxa is met with in many genera of fleas, but not in

Ceratophyllus proper, not in Neopsylla, Ctenophthalmus, and Paleo-

psylla, except the two new Neopsylla herve described. A survey

of the genera which bear coxal spines renders it evident that such

spines have been acquired independently in many instances and do

not necessarily indicate close attinity. A species with spines may

be more nearly related to one without them than to another which

also bears coxal spines.

Five lateral bristles on the fifth tarsal segments are an ancestral

character for fleas, the segment with more than five such bristles,

or with the proximal pair shifted on to the ventral surface, or with

less than five pairs being more recent modifications, The five

airs are normal tor Ceratophyllus, while the species of Veopsylla,

Paleopsylla, Ctenophthalmus, and Amphipsylla have either five

pairs in the hind tarsus with the first pair placed in between the

second, or possess only four pairs. The present new species, there-

fore, connects that group of genera with normal Ceratophyllus.

But the interest of the species does not end there. We find that

the fifth tarsal segment of one hind leg has in both our females

on one side four and on the other five bristles, while the segment

has five on both sides in the other hind leg. In two of three

males, both hind legs have five bristles on both sides of the fifth

tarsal segment, whereas ina third male this segment has only fou

pairs of bristles. In all three instances where the number of

bristles is reduced, it is the proximal pair, or a bristle of the

proximal pair, which is absent. This appears to be decisive evi-

dence that also in the allied species which have only four pairs of

lateral bristles on the fifth hind-tarsal segment it is the first pair

which has been lost.

Head.— Evenly rounded, the frontal outline vertical in the ¢,

more slanting in the 9. Hye indicated by a narrow oblique bar.

Two genal spines, which cross each other as in many of the species

of this relationship, the outer spine being short and broad and the

lower one long and pointed. The genal lobe itself is narrower at

its base than in the centre, being sole-shaped. The frontal

SOME NEW CHINESE FLEAS. 389-

tubercle is small. ‘The frons bears two rows of bristles, the ante-

vior row containing in the ¢ 6 to 9 and in the @ 5 to 6 bristles,

the posterior row 3 long ones, in between which there are a few

minute hairs. The occiput has also two rows, an anterior one of

5 or 6 and a posterior one of 7 on each side, there being along

the antennal groove a number of small hairs, in the ¢ an irregular

row of about 18, and in the @ a patch of about 6 below the

last bristle of the posterior row and two or three hairs further

forward. The proportional measurements of the segments of the

maxillary palpus are 12, 10, 8, 16. The maxilla is long, reaching

to the apex of the third or base of the fourth segment of the

rostrum. The latter is one-third to one-fourth short of the apex

of the fore coxa; the fifth segment is more than twice the length

of the fourth. The antennal groove is continued to the vertex in

both sexes. The second segment of the antenna bears in the 9

three bristles, which extend beyond the apex of the club.

Thorax.—The pronotum bears a comb of 16 spines and a row of

15 or 16 bristles on the two sides together. There are on the

mesonotum a postmedian row of 14 long bristles, an anterior row

(sometimes double laterally) of 17 to 19, and a basal row of rather

long thin hairs, a few additional, short, dorsal bristles being placed

in the ¢ in front of the anterior row; before the apex of the

mesonotum there are 6 bristle-like spines on the underside. The

sides of the mesosternite bear 9 bristles. The metatergite has a

posterior row of 14 or 15 bristles, and an anterior row of 12 or 13.

The metepisternum bears 3 or 4 bristles, and the metepimerum

12 to 14 (4 to 6, 5, 2 or 3).

Abdomen.—The tergites I to VII bear each two rows of bristles,

the posterior row containing 18 ; the anterior row is reduced on the

posterior segments of the ¢ to a few bristles. The stigma stands

within the posterior row between the first and second bristle.

The basal sternite has the usual pair of ventral bristles and on

each lateral surface 2 or 3 more bristles. The sternites III to VIL

bear a row of 6 to 8 bristles in the ¢ and 10 in the 9, there being

also several (about 6 to 8) small bristles in front of the row. On

the seventh segment the row contains 14 to 16 bristles in the 9?

and 8in the ¢. Three antepygidial bristles,in ¢ the central one

about three times the length of the others, in the 2 not quite thrice

the length of the upper one and less than twice the length of the

lower one.

Legs.—The hind coxa bears a patch of about 18 spines on the

inner surface. The hind femur has two subapical ventral bristles

on the outside and one on the inside, there being also one small

lateral bristle on the inside at the basal fourth. The outer dorsal

bristles on the mid and hind tibie are strong and all of nearly

even size, there being 10 or 11 such bristles inclusive of the apical

ones, and four long dorsal bristles on the inner side of the edge.

The lateral bristles are 5 or 6 in number and form onerow. The

bristles at the hinder side of the first mid- and hind-tarsal segments

are likewise strong and not quite so long as the corresponding

Proc. Zoou. Soc.—1911, No. XX VI. 26

390 DR. K. JORDAN AND HON. N. C. ROTHSCHILD ON

bristles of the tibia, the first mid-tarsal segment bearing 4 or 5

such bristles and the hind-tarsal one 5 or 6. The bristles on the

anterior side of the first tarsal segments are much less numerous.

The two longest apical bristles of the hind tibia reach far beyond

the apex of the first tarsal joint, the longest of this segment

extends beyond the tip of the second segment, the longest of the

second segment to the middle of the fourth. The proportions of

the tarsal segments are :—

Horeltarsusi ic sll lO) NOs) 1S oli: mormon ali emo

Mid tarsus: @. 22, (8,135, 9) 19): Om 23522510, oO te

Hand tarsus: 6. 33,20, 08, 12) 195 Ora 2 a2 2 to 2.

The fifth segment has in all the tarsi five pairs of lateral

bristles, occasionally the basal pair or one bristle of it being

absent from the hind tarsus.

Text-

fi g. 121.

ae IXst.

o ; ee ee

= = = Mills

\ At

Text-

fig. 122.

Text-fig. 121.—Clasping organs of Neopsylla aliena 6.

Text-fig. 122.—Abdominal segments VII & VIII and receptaculum seminis of

Neopsylla aliena ¢.

Modified segments.— 3. The eighth sternite is broad, with

the apex almost evenly rounded in side view. It bears ventraily

on each side 14 to 16 bristles, of which two of the distal ones are

very long. The main body of the clasper (Cl) is almost square

(text-fig. 121). Its upper distal corner is rounded off and not

produced into a narrow process. There are a number of small

bristles at this corner, but no bristle near the insertion of the

SOME NEW CHINESE FLEAS. 391

movable process. The latter (F) is large, triangular, with the

distal side rounded and longest and the upper proximal side

shortest, the larger portion of the process lying from the point of

insertion downwards. The distal margin of this finger bears

numerous small bristles, which are nearly evenly distributed, one

below the apex being noticeable by its somewhat larger size.

The manubrium (M) is curved and widest in the centre, being

canoe-shaped. ‘The ninth sternite has a broad vertical arm, the

proximal portion of the horizontal arm also being broad. The

distal portion is at first narrow and then widens again, the apex

being curved upwards and ventrally furnished with seven pairs

of bristles (IX. st.)— 2. The seventh sternite (text-fig. 122)

is almost gradually narrowed in lateral view and divided by a

nearly central apical sinus into a rounded upper lobe and a broader

and less rounded ventral lobe. The eighth tergite bears 3 or

4 bristles above the stigma and several smaller ones proximally to

them. There are no bristles below the stigma, but the lower half

of the segment has 6 or 7 strong bristles along the edge, and | to

3 large ones on the side; proximally to these lateral bristles there

are about 6 to 8 smaller ones. At and near the apical edge, on

the inner surface, there are two rows of bristles, 4 or 5 at the edge

and 5 proximally to them. The upper apical angle of the segment

is produced into a prominent point. The eighth sternite bears no

bristles and ends on each side in a long slender point. The tail

of the receptaculum seminis is about twice as long as the head.

of the same.

Three males and two females from Yu-lin-fu, Shensi, 4000 ft.,

taken off Myospalax fontaniert.

STENOPONIA, gen. nov.

3 2. Similar to Hystrichopsylla Tasch. (1889), but differs in

very essential characters as follows :—

The labial palpus consists of only one segment instead of five

and does not extend much beyond the apex of the maxilla. The

genal process is narrow and short, and does not reach further

backwards than the uppermost spine of the genal comb. The

club of the antenna is short in both sexes, being but little longer

in the ¢ than in the 2. The sensory plate of the ninth

abdominal tergite is more convex than in Hystrichopsylla. The

first pair of bristles of the fifth tarsal segment is placed in between

the second pair in all the tarsi. The ? has only one receptaculum

seminis instead of two.

Type: tripectinata Tiraboschi (1902), described as Hystricho-

psylla.

13. STENOPONIA CG@LEsTIS, sp.n. (Text-figs. 123, 124.)

Q. Similar to S. tripectinata Tirab. (1902), but at once

distinguished by the different shape of the head. In tripectinata

the distance from the frontal corner to the first spine of the genal

26*

392 ON SOME NEW CHINESE FLEAS.

comb is about the same as the distance from this spine to the

last spine of the comb, whereas in the new species the former

distance is less than half the latter.

Text-fig. 123. Text-fig, 124,

oe es

Osi S Ie

; ae ie

an > \

: NS Lo SS sz = =a

aS, ‘ey ells INA tile

x =. oy i

KK lst

SS ad Nish

Text-fig. 123.—Head of Stenoponia ceelestis 2 *.

Text-fig. 124.—Abdominal segments VII & VIII of S. celestis 2.

Head.—The genal process, which is visible above the last spine

(text-fig. 123), is much broader than in tripectinata, and the

rostrum is longer than in that species. The second segment of

the antenna has two rows of bristles in both species. The genal

comb consists of nine spines.

Thorax.— The thorax is similar to that of tripectinata.

The pronotal comb consists of 35 spines.

Abdomen.—The tergites have only two rows of bristles each,

only segments I, II, and III having an incomplete third row,

while in é¢ripectinata all the tergites have 3 or 4 rows. There are

4 antepygidial bristles on each side. The comb of the first

tergite contains 31 spines on the two sides together.

Legs.—The bristles of the legs are shorter, especially those of

the hind tarsus, and also not so thin at the tip. The longest

apical bristle of the first hind-tarsal segment reaches only to the

subapical notch of the second segment, and the longest bristle of

the second segment scarcely extends to the apex of the third.

Modified segments.— 9. As these segments are represented

by text-fig. 124, we need only point out some characters in

which they differ from the segments of tripectinata. The lobe

* The lateral pale dot of the occiput 1s omitted.

ON THE ANATOMY OF FROGS OF THE GENUS MEGALOPHRYS. 393

‘below the sinus of the seventh sternite (VII. st.) is narrower

in tripectinata, and the lobe above the sinus longer than the lower

lobe, the sinus therefore being much deeper in that species.

The eighth tergite is divided by an apical incision into a broad

setose upper lobe and a narrower and naked lower lobe in both

species. The upper lobe, however, bears more bristles in tripec-

tinata both on the outer and inner surfaces, and the bristles

placed further proximad on the eighth tergite are also more

numerous in tripectinata. The eighth sternite, on the contrary,

has more bristles at the apex in the new species.

Length (mounted specimen, somewhat contracted) 3°5 mm.

One female from 23 miles $.E. of Ta-tsien-lu, 7500 ft., off

Scturotamias davidianus consobrinus M.-Edw.

20, Contributions to the Anatomy of the Anura. By Frank

EK. Bevparp, M A., F.R.S., F.Z8., Prosector to the

Society.

[Received December 28, 1910: Read March 7, 1911.)

(Text-figures 125-133.)

I. Some Noves upon tHe Froe JW/eG4aLoryrys

( LEPTOBRACHIUM) FEZ.

Of this species* living examples have been recently, and are

at the moment, exhibited in the Nociety’s Gardens. The Frog was

later described by Mr. G. A. Boulengert as of the genus Lepto-

brachium, but originally = referred to the genus Megalophrys,

to which all the Pelobatide belonging to the former genera

Megalophrys, Xenophrys, and Leptobrachium are now$ by him

referred. In dealing with certain points in the anatomy of

M. fee, I shall have occasion to refer to the mutual likenesses

and unlikenesses between this and other species of the family

to which I have already paid some attention ||.

The external characters have been so fully described by

Mr. Boulenger in the several papers quoted below, that little

remains to be said under this heading. There is, however, one

point to which I may refer. In dealing with Xenophrys monticola

and other forms, I have described 4 and figured a glandular patch

upon the thigh which is very characteristic of these Frogs. I can

find no trace of this structure in Megalophrys few ; and it is thus

* The specimens were, as I understand, identified by Mr. G. A. Boulenger.

t Ann. Mus. Civ. Genova, vol. vii. 1889, p. 750.

t Ibid. vol. iv. 1887, p. 512. For other references see Mr. Boulenger’s paper in

ie P.Z.3. quoted below.

§ P.Z.S. 1908, p. 407.

|| P.Z.S. 1907, p. 324, and ibid. 1907, p. 871. The latter paper deals with

Megalophrys montana, “ Xenophrys monticola,” “ Leptobrachium hasseltii,” aud,

incidentally, with Megalophrys nasuta, which is more fully described in the former

paper.

P.Z.S. 1907, p. 879, text-fig. 2

394 MR. F. E. BEDDARD ON THE ANATOMY

evidently not a distinguishing feature of the genus, as opposed, to }

Pelobates, as I had been disposed to think. 4

I now direct attention to cther points in the anatomy of this

species, and more especially to those which are already known to

be of systematic importance. This, indeed, has been my object

rather than to attempt a more comprehensive anatomical account.

$ Sternum.

In considering the mutual affinities of four species of Pelo-

batidz described by me in a former paper *, I had laid some stress

Text-fig. 125,

Sk. -

Abd. P.

Ventral view of antevior part of the body of Megalophrys fee partially dissected,

Sk. Skin reflected. Abd.P. Pectoralis abdominalis. O. Omosternum.

The cartilaginous right overlapping epicoracoid is dotted; the bony coracoid

and shaft of sternum are marked by short lines.

upon the proportions visible in the total length of the body on the

one hand, and of the sternum and sternal region on the other.

* P.Z.S. 1907, p. 891.

OF FROGS OF THE GENUS MEGALOPHRYS. 395

The measurements which I have just made upon Megalophrys

jee confirm the value of the results which I obtained from the

former series of measurements. Of Megalophrys fee I ascer-

tained the following measurements :—length of body from snout

to cloacal aperture 95 mm. ; length of sternal region from anterior

Text-fig, 126.

Sternum of Megalophrys fee from the ventral surface.

O. Omosternum, X. Xiphisternum.

Between the omosternum and the shaft of the sternum is seen the right,

overlapping, epicoracoid.

end of omosternum to posterior end of sternum 30 mm.; length

of sternum proper 17 mm. Considering the body-length as 10,

the other measurements will be respectively 3°2 and 1:8. The

proportions therefore are, it will be observed, exactly as in

‘the species Leptobrachium hasseltti. In a second specimen the

396 MR. F. E. BEDDARD ON THE ANATOMY

proportions were the same, the Frog being about 1°5-2 mm. longer

with a corresponding increase of length of from 5-1 mm. in the

sternal measurements.

It is, as is well known, the general rule among the Arciferous

Batrachia Salientia that the right epicoracoidal cartilage should

overlap the left ventrally. This was the case in three out of the

four examples of Megalophrys fee which I have had the oppor-

tunity of examining (see text-fig. 125, p. 394). In the fourth,

however, the converse condition occurred. It may be that this

individual is of a different species ; and yet I cannot find from

a study of Mr. Boulenger’s table of classification # any grounds

for identifying this individual with other species described by

him. It is, however, rather different in appearance from the

other Frogs ; its build is more slender, the pelvic width is less,

and the dorsal depression is much deeper and with a more arched

upper bordering ridge. Other depressions upon the head are also

more marked, I could not, however, detect any other external

differences, All the four specimens, I should observe, are males.

In view of the fact that the omosternum is already known to

differ among the species of Megalophrys (contrast, for example,

M. montana and M. nasuta), it is important to note the condition

in Megalophrys few. In this species (see text-fig. 126) it is, in

fact, quite as well-developed as in Megalophrys nasuta, from

which, however, it differs in various details. Its total length is

8 mm., which represents a size proportionately equal to, or not

very different from, that of Wegalophrys nasuta. The epicoracoids

are firmly attached to the basal portion of the omosternum, which

shows a trace of its double origin (?from the two epicoracoids) in

that it is divided longitudinally by a white fibrous seam. ‘This

part of the omosternum is, in fact, precisely as is figured by

W. K. Parker t in Pseudis. In addition to this there is a

distal piece which is single and not divided longitudinally, but

which is divided off by a seam from the basal portion of the

omosternum. This piece is cheese-cutter-shaped, as shown in

text-fig. 126,

$ The Hyoid and its Musculature.

The hyoid cartilages and bones, as I have already pointed out,

present some differences among the various species of Megalophrys

(s.0.). I have therefore attempted a careful study of this portion

of the skeleton in Megalophrys fee. 'The basal cartilage of the

hyoid has the usual broad form found in the allies of this- species ;

of this the processus anteriores are bowed inwards anteriorly

almost to meet in front of the exit from the hyoid region of the

hyoglossal muscle. ‘This condition of the processes in question

has apparently, as I have already pointed out, some systematic

importance. The hyoglossal sinus, thus nearly converted into a

* P.Z.S. 1908, p. 416. i

+ “A Mor Staph of the Shoulder-Girdle,’ Rae Soc. Publ. 1868, pl. vi: fig. Yio

OF FROGS OF THE GENUS MEGALOPURYS. 397

foramen, is of much greater diameter than the hyoglossal muscle

which passes through it. The imeonvenience which might thus

result is obviated by the existence posteriorly of a tough trans-

lucent membrane, which largely occupies the hinder part of the

sinus by stretching across it. It has a clean semicircular edge in

front over which the muscle plays. JI have been unable to find

any anterior cornua of the hyoid, which appears to be absent,

at any rate asa discrete cartilage, in this group of Frogs. But

the lateral foramina are present and correspond, as I imagine, to

similar foramina in Pelodytes punctatus, the development of which

has been worked out by Ridewood*. In Megalophrys fee they

transmit in the same way the glossopharyngeal nerves, than which

they are only just larger. The strongly ossified thyrohyals have

the usual elongated hourglass shape. The distal castilaginous

epiphysis of each of these bones has very much the shape and

direction that it has in Xenophrys monticola, to my figure of

which I refer below +.

The musculature of the hyoid does not appear to me to present

any particular features of interest as compared with allied forms.

The hyoglossus did not show the twisted rope-like strands that I

have observed and described in allied forms. Each muscle arose

solely from the inner and lower border of the thyrohyal bone, and

not at all from the flat surface of that bone.

§ Alimentary Canal.

The accompanying drawing (text-fig. 127, p. 398) of the ali-

mentary canal of Megalophrys fee may be compared with that of

Breviceps upon a later page (p. 407). The stomach of the Mega-

lophrys is longitudinally ridged by thick ridges, of which there

are eight in the middle of that organ. Of these only three survive

until the opening of the stomach into the duodenum ; the others

die away and cease to be ridges. The intestinal tract measures

104 mm., and is to be divided into a very short duodenum, a

wider ensuing region, and then a narrower ileum which opens

into the large intestine. ‘The regions are, in fact, quite as in

Lreviceps. ‘The length of the three different regions of the small

intestine are 6, 42, and 57 mm. respectively. Although it will be

observed that these measurements do not tally exactly with the

total length of the small intestine, it will be noted that they are

only divergent by 1mm. As the measurements of the different

regions of this gut were made quite independently and without

any concurrent reference to the total length of the small intestine,

I leave them at the figures which I have given. It will be observed

that these proportions are very different from those exhibited by

Breviceps. ‘There are also certain differences in the ridges which

* “On the Structure and Development of the Hypobranchial Sl:eleton of the

Parsley Frog (Pelodytes punctatus),” P. Z.S. 1897, p. 577.

+ P.Z.S. 1907, p. 398, text-fig. 238.

398 MR. F. E. BEDDARD ON THE ANATOMY

Text-fig. 127.

are RST tt ah eee ee

raven

}

The alimentary tract of Megalophrys fee from the middle of the stomach to the

middle of the colon laid open and divided into two portions, an anterior (to the

left) and a posterior (to the right of the figure).

St. Stomach. Co. Colon.

OF FROGS OF THE GENUS MEGALOPHRYS. 399

line the gut. But the general plan is the same. The first region,

which is so very short, is marked off from the stomach by a sudden

diminution in the thickness of its walls. The lining-membrane

is at first smooth and is later covered by ridges forming a reti-

culum. These gradually become the circular folds like those of

Breviceps ; but the region of the gut where the circular folds occur

is very limited and not more than 10 mm. long. The terminal

and narrower portion of the gut is marked by about ten longi-

tudinal folds, the transition between which and the transverse

folds is a network.

These folds are at first permanent and cannot be removed by

stretching the gut; later they exist, but can be removed by

stretching. Finally, this region of the gut becomes smooth before

its opening into the large intestine. The latter is at first smooth

but later has longitudinally running folds.

The liver viewed from the ventral surface is seen to consist of

a larger left lobe and a much smaller right lobe, between which

the large gall-bladder is very obvious and extends back a good

way beyond the edge of the liver into the abdominal cavity.

Under the large left lobe lies a smaller lobe, which is of about the

same size as the right lobe and is continuous with it under the

pericardium and membrane of the gall-bladder.

Lying above the viscera the large subvertebral lymph-space is

very obvious. Its relations to the body-wall and to adjacent

viscera are as follows :—The sac is bifurcate in form and extends

forward beyond the anterior end of the kidney, but not so far as

to the anterior end of the ileum. The two bifurcations of the

sac lie on either side of the ileum, and the sac therefore does not

extend so far forward where it lies upon the iliac musculature as

it does right and left of this region. It does not extend ventrally

to the kidney, but separates this viscus dorsally from the dorsal

body-wall. Posteriorly it overlies the bladder, which is firmly

attached to its wall.

§ Hsophageal Muscle.

This visceral skeletal muscle is so important and characteristic

in Megalophrys and its allies that its consideration demands a

section to itself. It is rather thin and does not overlap the

kidney, which lies entirely posteriorly to it. It does, however,

overlap the anterior part of the ileum. I have figured the muscle

in Megalophrys montana * as extending far back, in fact so that

it overlaps the sacral vertebra transverse process at its compara-

tively narrow origin from the centrum. This was not the case

with Megalophrys jee, where the muscle is therefore less extensive.

It is, however, large when compared with that of Frogs of other

families. It appears to differ but little in details from the

muscle of Megalophrys nasutat.

* P.Z.S. 1907, p. 886, text-fig. 233, m.

+ Loe. cit. p. 876, text-fig. 229, 6.

400 MR. F. E. BEDDARD ON THE ANATOMY

§ The Larynx.

I do not here figure the larynx of Megalophrys fee because it

agrees substantially with that of other Oriental Pelobatide figured

elsewhere by myself *. There are, however, as I have shown in the

memoir referred to, at least specific differences among the Pelo-

batide. I find that Megalophrys few agrees very closely with

Nenophrys monticola and thus difters from Leptobrachium hasselti.

The hypopharyngeal processes of the cricoid cartilages are well

developed and remain separate, 7. e. are not united into a single

bar. <A delicate semicircular br ‘onchial cartilage is quite obvious

attached or close to the anterior extremity of the cricoid cartilage.

§ Muscles of Ventral Surface.

In removing the skin of the ventral surface in the pectoral

and cephalic region the septa bounding the lymph-spaces in that

region come into view. I have already figured these in Xenophrys

monticola y, and the corresponding septa in Megalophrys fee

appear at first sight to be much the same. ‘The posterior of the

two is rather more chevron-shaped in the line of its attach-

ment perhaps; but this may be a matter of varying tension. On

the other hand, the same chevron line occurs in Leptobrachium

hasseltwit. In Megalophrys fee the line of insertion of the wall of

the subcutaneous lymph-sac does not extend obliquely downwards

on to the pectoralis abdominalis as in Leptobrachium. There

appears to be no invasion of this membrane by muscular fibres, and

in this the present species agrees with the two just referred to.

The anterior wall of the pectoral or thoracic lymph-sac has

been figured by myself as a single wall in the two species which

are alyore referred to, and as arising from the edge (posterior) of

the submaxillaris muscle. I find “precisely the same origin in

Meg galophr ys fee; but there is in addition another membrane

rising from the muscular surface behind this line which joins the

first-named to be inserted in common with it on to the skin.

This tent-like arrangement leads to the formation of an additional

sac, which is obviously triangular in section. I may, perhaps,

have missed something of the same kind in the other Megalo-

phryid Frogs which I have examined, Examined microscopically,

‘the two walls of membrane showed nothing but white fibrous

connective-tissue; there was no trace whatever of any muscular

invasion, and, indeed, the edge of the submaxillaris could be

plainly seen to give off no fines into the membrane. Inasmuch

as I observed this double wall in four examples of the Frog, there

‘can, I think, be little doubt of its being the normal arrangement.

A further exploration of this cavity shows that it is not a

lymph-sac at all; it is the gular sac which is thus intruded

* P. Z.S. 1907, p. 898, text-fig. 238, p. 899, text- fig. 239, p. 902, text-fig. 240.

+ P.Z.S. 1907, p. 882, text-fie. 231.

t Loc. cit. p. 884, text-fig. 232.

OF FROGS OF THE GENUS MEGALOPHRYS. 401

between two subcutaneous lymph-sacs. At each corner near to the

angle of the jaw is the orifice into the mouth-cavity, through which

a probe was, of course, passed to show that the aperture did actually

communicate with the mouth-cavity. On opening the mouth

the orifices in question were very plainly visible posteriorly. It

will be noted that the gular sacs in this Frog, instead of being

separate sacs as in /tana, are entirely confluent and form one

median ventral sac, which is shown in the accompanying text-

figure (text-fig. 128), as in Rhinoderma darwini*. There is no

trace of a division in the middle line of the body between the

right and left sacs; they obviously form a continuous single

narrow sac lying just behind the subhyoideus muscle opening at

Text-fig. 128.

Gular sac (S.) of Megalophrys fee.

On each side the dark shaded orifice into the buccal cavity is seen.

either extremity into the mouth-cavity. The walls of the gular

sacare rather extensive ; but, as has been already mentioned, they

are inserted in common on to the skin. This obviously would

hinder any very great dilatation of the sacs. Furthermore, they

could not extend in a dilated condition very far back along the

body, for the septum between the lymph-sacs in the pectoral region

would clearly prevent this. In Lhinoderma, on the other hand,

“the main portion of the sac hangs free” f.

As to the musculature proper, the rectus abdominis shows no

definite traces that were at all apparent to me of itnseriptiones

tendinece, which seems to be rather an important difference from

it nearest allies.

* Cf. Howes, “Notes on the Gular Brood-pouch of Rhinoderma darwini,” P.Z.S.

1888, p. 231.

+ Howes, loc. cit. p. 234.

402 MR. F. E. BEDDARD ON THE ANATOMY

The pectoralis abdominalis is not very extensive; it reaches

back for about one-third of the distance between the end of the

sternum and the symphysis pubis.

The musculature in the sternal region (see text-fig. 125, p. 394)

does not entirely cover the bones and cartilages of the shoulder-

girdle. The right epicoracoidal cartilage, which greatly overlaps

ventrally the corresponding cartilage of the left side, is for

the greater part bare of muscle. A large portion of the right

coracoid bone and the triangular bony base of the sternum

were also uncovered by muscle. In a second specimen, however,

the pectoralis did extend over the triangular basal region of the

sternum.

The rectus abdominis continues forward anteriorly after the

origin of the pectoralis abdominalis and lies to the side of the

sternum, separated from it, however, by a much narrower muscle

which arises from the shaft of the sternum, and which I shall

again refer to as a portion of the sternohyoideus. At the

anterior end of the dagger-shaped sternum a portion of the

rectus abdominis is inserted upon the lateral transverse extension

of the ‘“‘handle” of the “dagger.” The rest passes onwards

anteriorly and becomes a part of the sternohyoideus. I did not

observe in the present species a tendinous connection of the rectus

with the edge of the coracoid such as that figured in Xeno-

phrys monticola *. Nor did I note any fibrous lateral expansion

of the sternum, such as occurs in the species just mentioned,

covering over the innermost section of the sternohyoideus muscle.

If such a membrane were present it must have been excessively

thin and delicate to have escaped observation.

I have mentioned that in Xenophrys no superficial sternoradialis

is visible; in the present species it is only superficial—and then

covered by fascia—for a very short distance after its origin from

the omosternum. It is thereafter covered by the pectoralis. The

strong tendon was followed to its insertion, and there is thus no

doubt about the presence of this important muscle in Megalophrys

fee. As the fascia in question is continuous with the pectoralis,

the sternoradialis may be said to be completely covered by that

muscle. The two (anterior and posterior) portions of the pectoralis

sternalis are very distinct at their origin, on the right side at any

rate, by reason of the exposed bony end of the coracoid which

separates them. The origin of the pectoralis sternalis posterior

extends about halfway down the bony shaft of the sternum, but

appears to vary in individuals, in the extent of its attachment to

the sternum, as the text-figure (text-fig. 125) referred to above

shows.

The throat musculature of Wegalophrys few is exhibited in the

accompanying drawing (text-fig. 129). There are no prominent

differences from the conditions of the corresponding muscles

* P.Z.S. 1907, p. 893, text-fig. 236, a.

OF FROGS OF THE GENUS MEGALOPHRYS. 403

observable in other Megalophryide. The submentalis is large

and conspicuous anteriorly. In the middle of the area occupied

by the two succeeding muscles is a tendinous sheet quite similar

to that which I have figured in Leptobrachium hasseltti*. ‘This is

concerned with the subhyoideus as well as with the submaxillaris

muscle. The fibres of the latter radiate a good deal from the

central tendon—that is to say, they do not cross the jaw space

Text-fig. 129,

Ventral musculature of the throat of Megalophrys fee.

Sk. Skin reflected. A. Subhyoideus. B. Submaxillaris.

Anteriorly to the latter is the submentalis.

evenly and in parallel lines from side to side, being only interrupted

by the central tendon. The line of demarcation between the sub-

maxillaris and subhyoideus is very distinct indeed. The sub-

maxillaris, it should be remarked, completely covers over the

submentalis, though it thins off in this anterior region of the

throat.

§ Musculature of the Dorsal Surface.

The depressor mandibule seems to be very much as in Rana

and is a stout muscle firmly attached in front to the skull, where

it shows thicker in transverse section ; it completely covers the

scapula together with the latissimus dorsi, the two forming a con-

tinuous sheet of muscle. I have not mentioned the depressor

mandibule in my two former papers upon the anatomy of the

Pelobatide, and therefore take this opportunity of asserting its

presence in the genus Megalophrys.

* “Contributions to the Knowledge of the... Pelobatidew,” P.Z.S. 1907, p. 884,

text-fig. 232.

AY 4 MR. F. E. BEDDARD ON THE ANATOMY

Tl. Furtuer Nores upon tHE Genus BREVICEPS.

Nearly three years ago I communicated to the Society some

notes upon the anatomy of an African Frog belonging to the

genus Breviceps, which I identified with the species B. verrwcosus *.

I am now of opinion that three Frogs upon which I reported on

that occasion are not to be referred to this species, which I have

lately examined. The specimens of Breviceps verrucosus and of

B. gibbosus, which were exhibited at the Society’s Gardens some

years since, and of which several have been preserved in alcohol,

are distinctly different from those to which I had devoted my

attention in the paper already quoted. They are much more

spherical in outline, agreeing with a published figure of B. verru-

cosus ¢ and of B. mossambicust. ‘The examples formerly studied

by myself were much longer in proportion and of a squarer out-

line. Furthermore, the examples of B. verrucosus and BL. gibbosus

in the possession of the Society are more tubercular upon the

surface of the body. I am, however, quite unable to suggest the

species to which the specimens dissected by myself in 1908 are to

be referred. I cannot at any rate refer them to Mr. Boulenger’s

recently described Breviceps macrops §, for that species is abnormal

(for Breviceps, though more normal when compared to other Frogs)

by reason of the relatively large head and eyes. Breviceps pentheri

of Werner || is quite too small a species to be confused with that

to which I now refer, although the latter is considerably smaller

than either B. verrucosus or B. gibbosus. The latter are hard to

separate, as Mr. Boulenger has pointed out. A sixth species,

B. adspersus J, appears to me to be too briefly described to permit

of its identification with either the species with which I am now,

or that with which I was, concerned.

The present contribution to our knowledge of this genus

Breviceps is based upon the examination of four examples given

to me by Mr. Purcell, who was so good as to have them collected

in the Cape Colony. These seem to me to belong either to

B. gibbosus ov B. verrucosus, which I cannot differentiate to my

own satisfaction and which are in any case very closely allied.

Through the kindness of Mr. Purcell they were very well pre-

served for dissection purposes, and I am thus able to add some-

thing to the existing knowledge of Breviceps. I have already

mentioned a few external differences between the present species

and those which I dissected in 1907. Ishall point out in the

course of the following pages certain anatomical differences.

* Rapp. Arch. f. Nat. 1842. p. 291.

+ Steindachner, Reise der Novara, Amphibien, Wien, 1867, pl. v. fig. 3.

+ Peters, Reise nach Mossambique, Amphibien, Berlin, 1882, pl. xxv. fig. 2.

§ Ann. & Mag. Nat. Hist. xx. 1907, p. 46. Mr. Boulenger remarks that,

“Unlike B. gibbosus, B. macrops does not cover itself with a viscous secretion when

alarmed.” The species upon which I report inthe present paper showed in one indi-

vidual a thick mass of secretion on the body.

|| Zool. Anz. xxii. 1899, p. 116; the species is 15 mm. long.

@ Peters, Reise nach Mossambique, Amphibien, Berlin, 1882.

OF FROGS OF THE GENUS BREVICEPS. 405

between these members of the same genus. On the other hand,

I am able to confirm certain structural features of the genus as

formerly described by myself by finding an identical arrangement

in these examples of B. gibbosus.

§ Genito-urinary Organs.

The testes, fat-bodies, and kidneys of Breviceps gibbosus do not

altogether agree in their characters with my former description.

There is no doubt that the vasa efferentia of the testes are

numerous, as is the general rule among the Anura. This fact I

am clear about. So obvious are the several parallel sperm-ducts

issuing from each testis, that I cannot understand how I can

have been in error in describing only a single duct in the smaller

species of reviceps, if, that is to say, I was in error. Moreover,

in Breviceps gibbosus the ureter arises from the lower corner of

the kidney, and not from rather higher up as I have figured it in

Breviceps sp.* The two kidneys are nowhere fused together in

the middle line. It is worth recording that the fat-bodies have

but few finger-like processes; I found two and three in one

specimen and three and four in another. But these structures

are known to vary.

$ Respiratory Organs.

The lungs of Breviceps gibbosus present several features of

interest. Hach lung itself (in the contracted condition in which

it appears in the alcohol preserved specimens) is broader at the

base and narrows towards the free abdominal extremity. It is

important to notice that both extremities of the lung are free and

that the bronchus enters that organ at about the second third of

its entire length. The lung is thus not merely a sac dependent

from a rudimentary trachea. Furthermore, two bronchi are very

plainly differentiated. Hach is, in fact, about half as long as the

lung (contracted, of course) into which it opens. It is proportion-

ately wide and enters the lung at right angles to the long axis of

the latter viscus. Its walls are membranous and translucent and

T could detect no cartilages. The pulmonary ligament fixing the

lung to the dorsal middle line extends along the whole of the

bronchus and just on to the lung.

If I have not in any way misread the conditions which obtain

in the species of Lreviceps dissected three years since, that

species shows considerable differences from Sreviceps gibbosus in

the relations of the wsophago-pulmonary muscle (“ diaphragm”).

There are, however, also points of agreement between the two sets

of individuals. For instance, the muscle arises in both in the

same way from a vertebral transverse process nearly in common

with the transverso-scapularis. Furthermore, in both cases the

* P. Z.S. 1908, p. 38, text-fig. 13.

+ Loe. cit. p. 27, and text-fig. 7, p. 28.

Proc. Zoou. Soc.—1911, No. XX VII. 27

406 MR, F, E, BEDDARD ON THE ANATOMY

c@sophago-pulmonary muscle is closely associated in function

with the hindermost of the petrohyoid. muscles,

Here, however, the principal resemblances end and there remain

certain important differences,

The most important of these points of difference is that the

muscle which we are discussing has no relations whatever,

except those of contiguity, to the respiratory organs. I believe

that I can assert this fact with confidence; but I have no reason

to doubt the essential accuracy of my description of the other

species of Breviceps. The difference in this particular may be

due to the extraordinary development of the bronchi in Breviceps

gibbosus. In this species, the insertions of the cesophageal muscle

and of the hindermost petrohyoideus are as follows :—Two muscles

are seen to be inserted on to the ventral surface of the esophagus,

fanning out somewhat at their insertion, which falls short of the

middle ventral line. Itis perfectly clear, when these muscles are

followed back, that they have no attachment (distal) other than

to the surface of the esophagus. The cesophageal muscle is the

larger of the two, and it partly forms an arch over the cesophagus

which is not attached to it; some of its fibres, that is to say, run

from one vertebral transverse process to the corresponding one

upon the opposite side of the body. The rest are inserted upon

the cesophagus ; but they pass underneath the fibres of the petro-

hyoideus and are inserted laterally to them. ‘The insertion of the

petrohyoideus upon the cesophagus appears to me to be quite as

extensive and important as is that of the cesophageal muscle

itself,

§ Alimentary Tract.

I have had represented in the accompanying text-figure

(text-fig. 130) the intestinal tract of an example of Breviceps

gibbosus, which has been opened up for the greater part of its

length. The general aspect of the intestine of this species 1s very

like that of the species which I formerly described *, and it

can be divided into precisely the same regions. In the present

species the duodenum measured 7 mm, in length ; the ensuing

wide region of the intestine was double this length, viz. 14 mm.;

the long narrow terminal region, which opens into the colon, was

again double the length of the preceding region and measured

29mm. The short duodenum shows externally a division into

small and approximately equal rounded areas, This appearance

was also seen internally where the folds of mucous membrane

constitute a network.

In the wide region of this gut the folds are circular and pre-

cisely as I have described them in the other species. In the

terminal region of the small intestine the folds are entirely

longitudinal, and there is a transitional area which is also

indicated in the accompanying text-figure.

* P. Z.S, 1908, p. 32, text-fig. 10.

OF FROGS OF THE GENUS BREVICEPS. 407.

The stomach has much the same character as in the other

species ; but the contents were quite different. In the stomach

described here were the remains of a largish beetle and an entire

and quite large beetle larva and some other insects,

Text-fig. 130.

Co.

Alimentary tract of Breviceps gibbosus with the greater part of the small

intestine laid open.

A-D. Portions’ of the different regions of the intestine more highly magnified.

Co, Colon, cut short. H. Heart, Z. Liver. St, Stomach.

The text-figure to which I have referred in describing the

intestinal canal (text-fig. 130) also shows the liver, which is a

little different from that of the other species of Areviceps whose

anatomy has been deseribed by myself. The heart is not at all

A08 MR. F. E. BEDDARD ON THE ANATOMY

covered by the liver in the species Greviceps gibbosus. ‘There is,

however, as I have illustrated by figures*, some individual

difference in the liver in this genus.

§ On certain Characters of the Skeleton.

I did not notice particularly the sternum of the other species of

Breviceps ; in B. gibbosus the form is not at all as it is figured by

Parker in his ‘ Monograph of the Shoulder-Girdle’‘?, nor is it

like the woodeut given by Boulenger in the ‘ Catalogue’=. The

cartilaginous plate is much more important than would be

gathered from those figures, and expands laterally into a curved

and thinnish process. It is, in fact, very like the sternum of

Discoglossus pictus as figured by Boulenger §.

Two other features in the skeleton of this frog were commented

upon by myself in the earlier account of the anatomy of this

genus. I naturally endeavoured to ascertain if those peculiarities

were to be found in Breviceps gibbosus, and I find that the

species which forms the subject of the present notes has the plate

of cartilage lying upon the expanded sacral transverse process,

and that the anterior cornua of the hyoid are perforated in

exactly the same fashion ||. I need not give a more prolonged

description of these peculiarities, which appear to be identical in

the two species.

§ On certain Muscles.

Many of the muscular peculiarities of the species described in

my earlier paper occur also in the species now under consideration.

It is, perhaps, impertant to enumerate such of these as I have

verified, in order to confirm them—for they are unusual—and

also in order to point out that they are apparently characteristic

of this genus or at least of more than one species. I shall not,

however, give a long description of them. In the first plaee, the

latissimus dorsi (see text-fig. 131), not a very large muscle, is

completely covered over by the double sheet of the obliquus.

There is also no trace of the posterior part of the depressor

mandibule. I did not note in my paper whether the cephalic

portion of this muscle was present. I find that it is present

in B. gibbosus. In the present specimens also the suprascapula

is largely exposed anteriorly when the skin is removed, on account

of the absence of the posterior part of the muscle just referred

to. It is covered, however, by a tough fascia. Furthermore,

the dorsal muscle of the suprascapula, the injraspinatus, is quite

visible without removing any other of the dorsal muscles.

The rectus abdominis is nearly as in Sreviceps sp., where

I have figured it. In B. gibbosus I could find one superficial

* P. Z.S. 1908, p. 30, text-fig. 8. + Pl. vii. fig. 9.

+ ‘Catalogue of Batrachia Salientia,’ 1882, p. 176.

§ ‘Tailless Batrachia of Europe,’ Ray Soc. Publ. 1897, vol. i. p. 40.

| Of. P.Z.S. 1908, p. 12, text-fig. 2.

OF FROGS OF THE GENUS BREVICEPS. 409

inscriptio tendinea *, behind the origin of the pectoralis abdomi-

nalis.. This muscle arose a little in front of the first and only

visible inscriptio tendinea.

The other abdominal muscles, q.s. emo-abdominal, hyo-abdoni-

nal, and obliquus internus, are quite as IT have described them in

my former paper, except that the hyo-abdominal seems to le

rather larger. This muscle passes under the lateral process of the

sternum, ¢. e. dorsally to it, and would thus appear to belong to

the obliquus internus sheet, and not, as I formerly suggested, to

the obliquus externus.

Text-fig. 131,

Cu.

Obs. int.

Breviceps gibbosus, from the dorsal surface, partly dissected.

Cu. Cutaneous muscle. Z.D. Latissimus dorsi. Obl.int. Obliquus internus.

Sc. Seapula. «. Slight bulge of dorsal musculature referred to in text.

The obliquus externus is rather different from that of the species

of my former description. The fascia dorsalis is quite obvious,

and the origin of the muscle is thus some way removed from the

* Loe. cit. p. 23, text-fig, 5

410 MR. F. E. BEDDARD ON THE ANATOMY

middle line of the back (see text-fig. 131). Its fibres run obliquely

backwards and are massed into coarse strands at the origin of the

muscle. The origin of this muscle extends further back than was

formerly described, and covers the obliquus internus until very

near to the end of the line of origin of the latter. The obliquus

internus only becomes superficial for a very short distance pos-

teriorly. These differences may be seen on a comparison of

Text-fig. 132.

An enlarged view of a portion of text-fig. 131, to show absence of diverticulum

of body-cavity overlying thigh,

C. Cutaneous muscle. C7. Cloacal aperture. 067.7. Obliquus externus.

Ob/.ii. Obliquus internus.

text-fig. 131, p. 409, of the present communication, with text-

fig. 4, p. 20, of my paper on the other Lreviceps. I am able to

confirm the statements about the latter species, so far as the

points at issue are concerned, by the examination of a specimen

given by me to the Royal College of Surgeons, and dissected by.

My. Burne.

The aforesaid figure (text-fig. 131), which is to be compared

OF FROGS OF THE GENUS BREVICEPS. 411

with text-fig. 4 of my previous paper upon this genus, not only

shows the differences in the oblique muscles already referred to,

but indicates the nearly total absence of anything comparable to

those structures which I identified in my former paper with very

much enlarged lymph hearts. There is, indeed, a slight bulge of the

abdominal wall posterior ly, but it does not extend over the dorsal

surface of the thigh; nor is this bulging at all detached from the

Text-fig. 135

Int. Ob/.

Gu:

Copy of a drawing by Mr. R. H. Burne, illustrative of certain parts in the

anatomy of Breviceps sp. referred to in the text.

Bl. Body-cavity. Cu. Cutaneous muscle. Jnt.06l. Obliquus internus. ZL. Lung.

Py. Pyriformis. S. Diverticulum of body-cavity overlying thigh.

The muscles of the thigh are indicated by dotted lines.

abdominal wall. It will be easily seen from an inspection of text-

figure 132 that an extension of the body-cavity on to the dorsal

surface of the thigh is impossible in the present species. For,

as will be seen in the last figure referred to, a ridge of cutaneous

muscle ties the commencement of the thigh to the skin and thus

412 ; MR. F, E. BEDDARD ON

effectually prevents any hernia-like outpushing of the abdominal

eavity backwards. 'Thereis thus an important difference between

the species with which I am now concerned and that which

formed the subject of my earlier paper.

With reference to the latter species, I am able, through the

kindness of Mr. R. H. Burne, to submit a drawing of a dissection

made by that anatomist on a specimen of the toad which

I presented to the Royal College of Surgeons. Having also seen

the actual dissection, I am obliged to admit that Mr. Burne has

proved that my former account of this region of the body was not

correct in every detail. Mr. Burne has ascertained and has

demonstrated to me that the structure which I termed a ‘“ lymph

heart” is a diverticulum of the body-cavity, lying, however, on the

leg, as is clearly shown in the accompanying drawing (text-

fig. 133, p. 411). This connection was so wide that the bladder

had floated into the diverticulum.

T feel convinced, however, that the communication was not so

wide in the larger female example dissected and figured by myself.

For the diverticulum was easily detached from the surrounding

structures, a fact which argues some independence, as it would

seem at least to show that the orifice into the general thoraco-

abdominal cavity can contract. Furthermore, the arrangement

of the oblique muscles in the region is rather different from what

may be seen anteriorly. Thus we have, certainly in this species,

a specialized portion of the thoraco-abdominal cavity (which is

recognisable, but much less prominent, in Breviceps gibbosus)

extending over the dorsal surface of the thigh. It is not,

however, I now admit, possible, in the present state of our know-

ledge, to speak of this as a “‘ lymph heart.” I propose, however, to

defer any further consideration of this subject until more facts

have been accumulated.

21. On the Spermatophores in Earthworms of the Genus

Pheretima (= Pericheta). By Frank E. Brepparp,

M.A., F.R.S., F.Z.8., Prosector to the Society.

[ Received December 28, 1910: Read March 7, 1911. ]

(Text-figures 134-136.)

I believe that there is no account of the spermatophores in

this genus of Karthworms; and at least there has not been, to my

knowledge, anything more than the briefest reference to their

occurrence. It might be expected from analogy that this genus,

like so many others, possessed this means of impregnation; but

I can recall no figures of such structures. Even supposing that

I have involuntarily ignored such an account, it is worth while

to add something more to the subject, which cannot be well

known.

SPERMATOPHORES IN EARTHWORMS. 413

I have lately examined, and shall shortly report upon, a collec-

tion of terrestrial Oligocheta from the Philippine Islands, the

opportunity of examining which LI owe to the kindness of the

Director of the Scientific Bureau of the Philippines. I found

these structures in two species contained in that collection, one

of which I propose to describe as a new species, and the other of

which is, as I believe, identical with Pheretima montana.

Tn the first-named species, which is very close to that which 1

call, in my revision of the genus*, P. cingulata, there are four

pais of spermathece. In all of ee. in individuals which I

examined for the present purpose, I found spermatophores, but

only one or two in each pouch. They are oval bodies with a long

spout-like projection, like a pear with a long stalk. It appeared

to me, on examining these mounted in glycerine entire, that the

prolongation did not open on to the exterior, and that the sperm

had therefore no obvious means of escaping from the case which

contains it. If my observations are correct, they do not, as will

appear shortly, imply anything anomalous; for the sperm has

another way of escape. In the species which I identify with

P. montana there is only a single pair of spermathece. But in

each spermatheca was a much larger number of spermatophores.

There is an obvious relation here between the total number of

spermatophores in the two species. In the former species, more-

over, the spermatophores had a longer spout-like projection than

in P. montana.

The accompanying figure (text-fig. 134, p. 414) illustrates a

spermatheca of P. montana, cut open and still containing two of

the spermatophores, while two others lie in the vicinity. There

are also three other spermatophores in this particular sperma-

theca, making a total of seven. In other cases I have noticed the

same, or nearly the same, number of these bodies. The figure

referred to shows the shape of the spermatophores quite well.

They vary slightly in size, beg not more than a millimetre in

length, and are spherical to oval with a narrow duct-like prolon-

gation. The shape, in fact, is quite consistent with the view that

they are spermatophores. They do not, however, agree absolutely

in their form with those of any Annelid which has been up to

the present described. In view of the considerable differences

which spermatophores show in different genera and families of

Oligocheta, this would hardly be expected. On the whole it

appears to me that they most resemble those of Pareudrilus,

described by myself? and Cognetti de Martiis <.

The Se matophores when viewed with the naked eye or with

a hand-lens havea glistening white appearance, which is explained

by their minute structure, as will be apparent later. J did not

* “« A Revision of the Genus Amyntas,” P.Z.S. 1900, p. 609.

+ “On a new Genus and two new Species of Earthworms, &c.,” P. Z.S. 1903,

Pp. 210.

{ “Contributo alla Conoscenza della fecondazione negli Oligocheti,’ Atti Acc.

Sci. Torino, xlv. 1910.

414 MR. F. E. BEDDARD ON

observe any case where the spermatophores were sticking on to

the surface of the body of the worm, as they do in the Lumbricide,

in Alma, in Bothrioneuron, and among the Leeches. The fact

that so many were crowded together in one spermatheca in the

species figured here seems to be some evidence in favour of

regarding the spermathece as wot being their place of origin.

There are also some other facts which favour the same negative

supposition. The size of these bodies is small; and it seems

ditticult to imagine that they could be moulded in the spermathece,

which are comparatively so large. It might be held, indeed, that

the narrow spout in which the spermatophore ends was moulded

by the compressed duct of the spermatheca; but this leaves the

larger body of the spermatophore unaccounted for.

Text-fig. 134.

Sp.

Spermatheca (C.p.) of Pheretima montana containing spermatophores (Sp.).

I believe that I showed some reason for regarding the

spermatophores of Denhamia austeni* as entirely formed in

the spermatheca and its appendage. But it is equally or perhaps

more certain that in other species of Oligochzta the spermato-

phores are moulded in the glands appended to the male ejaculatory

* “On the Spermatophores in the Genus Benhamia,” P. Z.S. 1901, p. 704

SPERMATOPHORES IN EARTHWORMS. 415

apparatus. Although one might suppose that in two genera, which

are probably to be regarded as nearly akin, the formation of the

spermatophores would be identical, there is in the Pheretima now

under consideration some positive evidence to show that the

spermatophores are formed in the terminal sac of the male

ejaculatory apparatus. In this species, as in other Pheretimas,

the spermiducal gland ends in a narrow muscular duet which

appears to open into a wide terminal bursa copulatrix. The

latter, of course, opens on to the eighteenth segment by a

conspicuous orifice. When sections are made through this

terminal apparatus it is. seen that the cavity of the bursa

copulatrix is by no means so large as the peripheral measure-

ments of the sac would lead one to imagine, but that the

greater part of its cavity is occupied by a large penis, which can

doubtless be protruded. ‘This penis, which is of a conical form,

is surrounded by the cavity of the bursa, which, however, lies

chiefly to one side, and there is in that region, therefore, a space

of moderate dimensions which is not far from the size of one of

the spermatophores. Furthermore, the cells which line this cavity,

except near to its external orifice, are tall, and the glandular-

looking cells stain badly, and thus appear to be full of

some hardly stainable secretion. The muscular duct leading

from the spermiducal gland is joined at about the end of the

first fourth of the penis by the sperm-duct (here single and

apparently not ciliated), and the conjoined duct opens not on

to the end of the penis, but rather to the side, 7. ¢. inside the

cavity of the bursa copulatrix, so that it might inject the

sperm into the cavity of the bursa. This sperm might then be

surrounded by an excretion of the glandular cells of the bursa,

and thus emerge a complete and fully charged spermatophore.

This is admittedly a mere suggestion, and is far from being a

conclusive statement. I have, however, no further evidence.

Reverting to the spermatophores themselves, they show, when

examined entire with a low power of the microscope, an opaque

appearance, less marked naturally at the free tail-lke termination.

The opacity is doubtless responsible for the white appearance of

the body, and is very different from what one would expect in a

spermatophore. Presumably with chitinous walls it would

be supposed that it would present a transparent, or at least

translucent, appearance when examined by transmitted light.

This is, however, not the case at all. Moreover the walls of “the

spermatophore have not a regular outline, but are roughened, as

if many minute particles were adherent to the outside. This is

not in any way different when the bodies are examined in

glycerine. The roughened and opaque exterior prevents a clear

view of the contained sperm, and, indeed, it would be impossible

to state from such an examination only that there was any sperm

within. Viewed in its entirety in glycerine not much more is

to be learnt about the spermatophore than is taught by an

examination of it as an opaque object with a lens. I have,

416 MR. F. E. BEDDARD ON

however, studied these bodies by the section method, which has

allowed me to describe a good many facts concerning their

structure, though something remains to be discovered, as will

in due course be pointed out. The wall of the spermatophore

is the first part of the structure which calls for attention. This

is of some thickness, as is shown in the accompanying figure (text-

fig. 135); but it is not thicker than that of other spermatophores.

Its structure, however, is remarkable, and in some respects

unique, at any rate at first sight. The minute structure of the

Text-fig. 135,

Longitudinal section through spermatophore of Pheretima montana.

S. Spermatozoa massed at apical end of case.

M. Mucous and granular substance filling up blind end of case.

wall is illustrated in the accompanying figure (text-fig. 136).

At first sight, the wall of the spermatophore suggests a living

tissue allied to connective tissue or even muscle. We recognise

many nuclei, which occur outside, inside, or within the wall itself.

These nuclei are not in any way degenerate structures, and their

varying position shows that they are not merely fragments ad-

herent to a sticky structureless wall. For they lie within as well as

on both sides of it. In addition to these cells there is a structure-

less substance which in parts has a fibrillar character. The likeness

to muscle is thereby much enhanced. It occurred to me at first

that the wall might be actually an adventitious sheath formed

by the tissues of the worm’s body, and comparable to the sheaths

found round foreign bodies when introduced from the outside, or

SPERMATOPHORES IN EARTHWORMS. 417

to the “ paruterine organs” of Tapeworms. It may be that this

interpretation is correct. But I do not myself hold this view

for other reasons, which will be explained. That portion of the

wall of the spermatophore which is lettered “Ch” in text-fig. 136

seems to me to give the clue to the real nature of the wall. It

will be noted that we have a thickish and structureless layer

which is surrounded on both sides by cells. The clear layer

is not much stained by the carmine, and is quite reasonably

to be regarded as the actual unaltered structureless wall of the

Text-fig. 136.

Transverse section through wall of spermatophore of Pheretima montana much

more highly magnified than in text-fig. 135.

Ch. Chitinous layer. NV. Nuclei of phagocytes (?).

spermatophore, probably of chitinous nature. Elsewhere the

same structureless substance is found which shows evidence of

being hollowed out. These facts thus briefly stated lead me to

the inference that we have here a spermatophore in which the

original chitinous wall is being gradually eaten away by leuco-

cytes. Ido not see any other view which fits in so well with both

fact and probability.

Indeed, the only other interpretation of the histological facts,

which is not very much borne out by the relations of the cells to the

membrane, is to suppose that the case of the spermatophore is

built round a mass of sperm lying in the spermatheea, a suggestion

which, apart from the actual facts, is not at all probable. I feel

418 MR, F. E. BEDDARD ON

fairly confident that the phagocyte theory is the correct one to

explain the anomalous appearances seen in the case of the sperma-

tophores of this Pheretima. As to the origin of the phagocytes,

I have no facts to offer. This is, however, not a weak point in

the argument that they are phagocytes. The cells are as cells

small; the nucleus is very large compared with its surrounding

rotoplasm, But the immense numbers of the cells makes up

for the lack of non-nuclear protoplasm, and the large nuclei are

an indication of their activity. The presence of these cells leads

to the rapid and dark staining of the spermatophores, which is

very noticeable, and not what would be expected in such a

structure, on the view that its walls were entirely chitinous. I

obtained the clue to the nature of the processes going on in the

spermatophores of this Oligochete when within the spermatheca

by a consideration of an important recently published paper by

Dr. Cognetti de Martiis. The author promises a further contri-

bution on the role played by phagocytes in the destruction of

superfluous spermatozoa; but this memoir I have not yet seen.

In the memoir which I have read and the title of which is quoted

below *, Dr. de Martiis observed that the spermatozoa leave the

large spermatophore, and suggests that the fibrils of the wall of

the latter are caused to diverge, and thus leave room for the exit

of the spermatozoa, by the action of certain cells against which

the end of the spermatophore is pressed, and which form part

of the lining epithelium of the spermathecal sac. A chemotactic

influence leads on the spermatozoa from cell to cell until they

reach the egg-sacs. He thinks that the weak and dying sperma-

tozoa are devoured by phagocytes, and has figured spermatozoa

in the interior of phagocytes. This, however, is a quite different

series of events from what takes place, as I believe, in Pheretima.

Here, I repeat, there is some evidence for regarding the phago-

eytosis of the walls of the spermatophore as a means of liberating

the enclosed sperm.

The contents of the spermatophore now demand attention.

The cavity of the spermatophore is not entirely filled with

spermatozoa, As will be seen in text-fig. 135, the mass of sperma-

tozoa is limited to the lower half of the spermatophore. It

occupies hardly half of the whole space available. The niass

of spermatozoa is strictly delimited above, and the line of demar-

cation is a regularly curved line. I think it possible to detect a

delicate membrane surrounding the mass of spermatozoa and to

be distinguished from the substance to be shortly described which

fills up the rest of the cavity of the spermatophore. The inclusion

of the spermatozoa in a case independent of the case of the

spermatophore itself, and lying within it, is a fact of likeness

to the spermatophores of Benhanuat, where something of the

same kind occurs. It is possible also that the delicate case of

the spermatozoa is fabricated in the appendix of the spermatheca.

% “Contributo alla Conoscenza della fecondazione negli Oligocheti,” Atti Acc.

Sci. Torino, xlv. 1910,

7, Bowes) 19018

SPERMATOPHORES IN EARTHWORMS, 419

But if this be so, the whole question of the place of formation

of the spermatophore will have to be reconsidered, I have, indeed,

no positive facts concerning the reasons for the rounded outline

of the mass of spermatozoa,

The arrangment of the spermatozoa requires some description.

They are not massed higgledy-piggledy with the heads and

the tails pointing in any direction; the arrangement is a

perfectly regular one, The heads of the spermatozoa are

all on that side of the sperm-mass which is nearest to the

“stalk” of the spermatophore. The upper part of the sperm-ball

is composed entirely of the tails, which are not disposed in

straight lines, but are waved and curved in every direction,

The heads radiate outwards in a quadrant or a little more

perhaps. The actual heads are in close contact with the walls

of the spermatophore, and some of them descend for a space

into the narrow duct of that organ. This rather looks as

if they were so disposed for easy egress at the time when the

sperm has to leave the spermatophore, which suggests that the

phagocytosis of the case is not a necessity for fertilisation. It

may be also that the spermatozoa escape, as Cognetti de Martiis

thought of the spermatozoa of Pareudrilus pallidus, through the

actual membrane of the spermatophore case, the action of the

phagocytes facilitating their egress by widening the strands of

the wall.

In any case the fan-like radiation of the heads of the spermatozoa

agrees with the idea that those which are more laterally placed,

and do not face the external spout-like orifice of the case, may

make their way out through the actual walls.

In addition to the spermatozoa the spermatophore contains a

granular mass which fills up the available space above the delicate

case containing the spermatozoa. This granular mass has the

appearace of broken-down cells. A comparison at once suggested

itself with the granular substance described by Whitman in the

spermatophores of Clepsine*. In the Leech, however, the

granular contents of the spermatophore lie near to the external

orifice of the spermatophore, and seem to clear the way from in

front of the subsequently issuing spermatozoa, or to perform

other functions which are discussed by Whitman. Like Whitman,

I was first disposed to regard the granular contents of the

spermatophore of Pheretima as being of a cellular nature. I

believe, however, that the substance is not composed of cells,

though probably of broken-down cells.

The position of this granular mass at the apex of the spermato-

phore suggests that it may be of mechanical assistance 7 in

expelling the sperm, supposing that the latter is usually expelled

through the mouth of the spermatophore, and not liberated by

* Journ. Morph. vol. iv. p. 361.

t+ See also Brumpt, “ Reproduction des Hirudinées,’ Mém. Soe. Zool. France,

1900, p. 286. But Kovalevsky (Comptes Rendus, vol. exxix. 1899, p, 261) did

find cells which he regarded as phagocytes to eat the spermatozoa.

420 MR. R. LYDEKKER ON

the eating away of the case. Furthermore, if the case be water-

tight, the presence of this possibly largely fluid mass may be

advantageous to the spermatozoa, especially if their sojourn in the

spermatophore be long. Its function may be to keep the sperm

moist and active. This is, however, mere supposition, and so

little is known about the processes of fertilisation in the Oligo-

cheeta that no safe guess can be hazarded. But it seems clear from

the large mass of granular substance that it plays some important

function in fertilisation.

22. A Rare Beaked Whale.

By R. LypEKKEr.

| Received December 19, 1910: Read March 7, 1911.}

(Text-figures 137-139.)

Some months ago—I believe early in the present year—a Beaked

Whale was stranded on the beach near Port Hlizabeth, which

fortunately came under the notice of Mr. F. W. FitzSimons, the

Director of the Museum in that city. Photographs were taken of

the specimen as it lay, and the skeleton was subsequently cleaned

and placed on exhibition in the Museum. As it lay, the specimen

measured 152 feet in length, from the tip of the muzzle to the end

Text-fig. 137.

Mesoplodon (Dioplodon) grayi as it lay on the beach.

The back-fin had been hacked by natives.

of the flukes. In colour it was jet-black all over; and the flukes

was remarkable on account of the posterior border being convex,

instead of deeply emarginate, as in ordinary cetaceans. The skull,

of which Mr. FitzSimons forwarded the two photographs herewith

reproduced, indicates that the specimen is referable to the genus

Mesoplodon (as commonly understood); this being manifest trom

A RARE BEAKED WHALE. 421

the presence of the single pair of large teeth just behind the hind

end of the symphysis of the lower jaw. These teeth are, however,

quite different in shape from those of Sowerby’s Beaked Whale (J.

bidens)—a species unknown in the Southern Seas ; and the whole

jaw is likewise different, as Mr. FitzSimons has pointed out, from

Text-fig. 138.

Palatal aspects of cranium and lower jaw of Mesoplodon (Dioplodon) grayi.

that of the South African Blainville’s Beaked Whale (JZ. densi-

rostris), in which the form of the teeth is also distinct. On the

other hand, in the great size of their basal portion and the upright

position of the terminal cap, the teeth agree exactly with those of

Proc. Zoou. Soc.—1911, No. XXVIII. 28

422 A RARE BEAKED WHALE.

the imperfect skull from New Zealand described and figured by Sir

William Flower, in the ‘Transactions of the Zoological Society,’

vol. x, p. 421, pl. Ixxii. fig. 3,.1878, as a new species, under the

name of Mesoplodon haasti, That so-called species has, however,

been identified by Dr. H. O. Forbes, in the Society’s ‘ Proceedings’

for 1893, p. 218, with the New Zealand species previously named

M. grayi, of which it represents.a very old individual, the func-

tionless small upper teeth found in younger specimens having been

lost. Assuming this identification to be correct, which I think is

probably the case, the Port Hlizabeth whale would appear to be

also a fully adult specimen of Mesoplodon grayi, and theretore of

great interest as showing the extension of the range of the species

to South Africa.

Text-fig. 139.

Lateral view of skull of Mesoplodon (Dioplodon) grayi.

But this is not all, for while, as already mentioned, the Port

Elizabeth Beaked Whale has the hind margin of the tail-fin convex,

in Sowerby’s Beaked Whale it is deeply emarginate, as is well shown

in De Blainville’s figure reproduced on page 255 of the ‘Study of

Mammals.’ <A feature similar to that found in the tail of the

Port Elizabeth specimen is stated to occur in a Beaked Whale

from Annisquam, Massachusetts, which Dr. F. W. True (Bull. U.S.

National Museum, no. 73, p. 10, 1910) refers to J. densirostris, a

near relative of M. grayi (with which J. australis is identical) ;

and it therefore seems that these two species form, at least, a

distinet sub-generic group, for which the name Dioplodon is

available.

I may add that I am indebted to my friend Mr. G. A.

Boulenger for handing me the original communication from

Mr. FitzSimons.

AGE-PHASES OF THE RORQUAL. 423

23. Age-Phases of the Rorqual.

By R. Lypexxker,.

[Received December 20, 1910: Read March 7, 1911.]

Some months ago I received a communication to the effect

that one of the captains at the Norwegian whaling-stations on

the Mayo coast was surprised to find that the true Rorquals of the

species commonly known to English naturalists as Lalenoptera

musculus found in the Irish seas differed from those he was

accustomed to take in Norwegian waters, not only in colour, but

likewise in their food. This species is stated in text-books to

feed largely on fish, especially herrings and pilchards; and those

taken in Norwegian waters by the captain above mentioned

appear to have been exclusively fish-eaters. On the other hand,

those taken on the Mayo coast in summer were feeding exclu-

sively on pelagic crustaceans. As to the nature of the colour-

differences between Norwegian and Irish examples, I was unable,

in spite of special inquiries, to obtain definite information.

I find, however, that Professor R. Collett, in a paper on Ru-

dolphi’s Rorqual, published in the Society's ‘ Proceedings’ for 1886,

states (p. 265) that in the True or Common Rorqual the whole of

the under surface of the flukes is white; this statement being

doubtless based on Norwegian specimens. On the other hand,

in the life-size model in the Natural History branch of the British

Museum of a specimen taken in the Moray Firth in the spring

of 1880 the under side of the flukes is coloured black. This

colouring is doubtless based on notes or a sketch taken when the

specimen was in the flesh; and, for what this is worth, its

correctness is confirmed by the fact that in the flukes itself,

which is preserved in the Museum, no difference in the colour of

the upper and lower surfaces is noticeable. In this connection it

is important to mention that specimens of this Rorqual have been

taken off Cornwall—in one instance in September—while feeding

on herrings and pilchards*.

This was the extent of my information on the subject till a

few days ago, when Dr. 8S. F. Harmer put into my hands some

notes on Norwegian Rorquals he had recently received from a

correspondent. From these notes, which are based on informa-

tion furnished by an experienced Norwegian whale-gunner, it

appears that the whalers of Finmarken recognise three varieties,

or phases, of True Rorqual—namely, a darker, a lighter, and a

yellowish. The dark phase is stated to be always met with in

company with shoals of herrings, upon which it doubtless feeds.

These Rorquals usually arrive off the Faroes in the middle of

June, coming from the south-west and proceeding in a north-

easterly direction ; the immigration lasting till about the middle

of July. Early in August the whales commence their return

journey southwards, when they keep principally to the south side

of the islands. | By the end of the same month nearly all have

* See Gray, Cat. Seals and Whales Brit. Mus. p. 149 (1866),

28%

4294. AGE-PHASES OF THE RORQUAL.

left the Faroes, the few that remain being restless and evidently

on the move.

The most important item in this statement is, however, that

these Rorquals arrive off Finmarken in two divisions, the first of

which is composed of members of the dark phase—which I presume

to be represented by the above-mentioned dark-tailed specimen.

These dark whales arrive early in March and advance in an

easterly direction, but do not, as a rule, go beyond the Varanger

Fiord, whence they return in the middle of April along the coast ;

the migration being completed by the end of the latter month.

The second division consists chiefly of the two lighter-coloured

phases, which I take to correspond with the light-tailed form

described by Prof. Collett. Nothing is said as to the nature of

their food; but it seems highly probable that it is this light-

coloured phase which is taken off Mayo feeding on shrimps, &e.

The members of this second division reach the Faroes in the first

half of June,,and are then met with about fifteen miles off Sord,

where they stay tillabout the beginning of July, when they move

eastwards as far as Baadsfiord, whence they finally disappear im

a north-easterly direction.

Schools composed chiefly of the light-coloured and yellow phases

reach Spitzbergen in the middle of June and move on towards the

north-east ; returning along the coasts of Spitzbergen from about

the end of July, and disappearing by the middle of August.

The existence of one dark and two light phases of this Rorqual

being thus clearly established, and it being also known that the

dark phase is the one associated with herring-shoals, it remains

to account for these differences. One explanation would be that

we have to do with distinet races; but although such an explan-

ation might be entertained if we had to do with two phases only,

it is difficult to accept when there are three. The alternative is

to regard the three colour-phases as representing differences in

age. Cetaceans of many kinds are known to show considerable

variation in colour according to age ; a notable instance being

afforded by coloured casts of two Norwegian examples of Sowerby’s

Beaked Whale (Mesoplodon bidens) exhibited in the whale-room at

the British Museum. In the smaller of these the belly is pinkish

yellow, whereas in the larger example it is as black as the back.

So different, indeed, are the two specimens, that I at one time con-

sidered they might be specifically distinct. I believe, however,

that Norwegian naturalists regard the black-bellied phase as the

fully adult condition of the white-bellied form; and if this be

correct, there is every reason for regarding the colour-phases of

the Rorqual as being likewise dependant upon age. Assuming the

colour-changes of the Beaked Whale to be paralleled in the Rorqual,

the dark phase of the latter should be the oldest ; and that this

is the case is indicated by the skeleton of the aforesaid specimen

in the British Museum, which is that of a very old individual, as

is shown by the complete welding of the epiphyses to the centra

of the vertebree. It may be also confirmed by the black-tailed

Rorquals being those which feed on herrings and pilchards.

LONGEVITY AND RELATIVE VIABILITY IN MAMMALS AND BIRDS.

425

24, On Longevity and Relative Viability in Mammals and

Birds ; with a Note on the Theory of Longevity. By

F.R.S:,

P. CuaumMers Mircueuyt, M.A., D.Sc.,

11 10),

Secretary to the Zoological Society of London.

[Received February 6, 1911

Tntroduction ..

Mammalia

Carnivora

Summary of Carnivora

Insectivora ( 67 examples)...

(292 HF, 1) ES

@330Gn ee ae JES

Summary of Rodentia............

( 12 examples)...

Proboscidea) (45 5) es

(1035 on Viens

Perissodactyla (46 _,, dese

Summary of Perissodactyla ...

Artiodactyla (989 examples)...

Summary of Artiodactyla......

Cetacea & Sirenia..................

Chiroptera

Rodentia

Hyraces

Ungulata

Edentata (123 examples)

Summary of Edentata............

Marsupialia (618 examples) ...

Summary of Marsupialia

Monotremata (4 examples)

Summary of Mammalia .........

Aves (12,977 examples)

Rassenesi( .o2240 seen) epee.

Summary of Passeres

Picariz (455 examples) .........

Summary of Picariz

Striges (432 examples)

Summary of Striges...............

Psittaci (1479 examples).........

Summary of Psittaci ............

Accipitres (893 examples)

Summary of Accipitres

(8110 ercvamleaye di

Quadrumana (8084 4, )...

Summary of Quadrumana ......

(1741 examples)...

: Read March 7, 1911.]

CONTENTS.

p. 425. ; Steganopodes (165 examples)... p.

p- 429. | Summary of Steganopodes......_ p.

p. 429. | Herodiones (480 examples)...... p.

p. 435. | Summary of Herodiones......... p

p. 486. | Odontoglosse (23 examples) ... p.

p. 443. | Palamedez (22 examples) ...... p

p- 444. | Anseres (1545 examples)......... Dp.

p. 444. | Summary of Anseres ............ p.

p-. 445. | Columbx (932 examples) ...... p-

p- 451. | Summary of Columbe............p.

p. 452. | Pterocletes (83 examples) ...... p-

p- 452. | Gallinze (1660 examples)......... p.

p. 453. | Summary of Galline ............ p

p. 458. | Hemipodi (15 examples) ...... p-

p. 454. | Fulicarize (214 examples) ...... p-

p. 434. | Summary of Fulicariz ......... p-

p. 461. | Alectorides (221 examples)...... p.

p. 462. | Summary of Alectorides......... p.

p. 462. | Limicolee (893 examples)......... p.

p. 463. | Summary of Limicole............ p

p. 464. | Gavize (241 examples)............ p.

p. 467. | Summary of Gavie............... p.

p. 467. | Tubinares (10 examples)......... p.

p. 467. | Pygopodes (86 examples) ...... p-

p- 470. | Impennes (62 examples) ......... p-

p. 470. | Crypturi (146 examples) ......... p.

p. 485. | Apteryges (22 examples) ...... p.

p. 487. | Casuarii (77 examples)............. p.

p. 491. | Struthiones (97 eee Ree p.

p. 492. | Summary of Aves.. te p.

p. 493. | General Summary wand Can:

p. 494. clusions . Dp.

p. 500. | Note on ihe cout ae

p. 501. IVORY snd hopuddsoodentaderents ) 10s

p. 505.

INTRODUCTION.

Animals are mortal, and we know that death comes to them

after durations of life that differ much in different individuals

and species.

Unfortunately, except in the rarest cases, it is

impossible to obtain information as to the duration of life in wild

animals.

Enquiry into the frequency of death in domesticated

426 DR. P. CHALMERS MILCHELL ON LONGEVITY AND

animals, or in animals in captivity, is therefore of scientific and

practical interest. From the scientific point of view, it is the

only mode of getting any kind of information as to constitutional

differences in longevity and viability. From the practical point

of view, it is the only way of judging of the effect on duration of

life of the different kinds of environment to which animals may be

subjected. There exists as yet very little recorded information,

and I hope that this memoir may at least stimulate enquiry and

make possible the collection and comparison of data.

The normal annual death-rate of the human inhabitants of

London is about 14 or 15 per thousand, or 1°5 per cent. per

annum, that is tosay for every two hundred inhabitants there are

about three deaths inthe year. There is no census of the sparrow

-population of London, but the numbers of these hardy birds do

not appear to increase or decrease in any marked way. Now

a pair of sparrows may rear two or three clutches in the course of

the year, each clutch having five or six eggs. If every pair of

sparrows alive in London at the beginning of the breeding-season

displayed maximum fertility, a death-rate of about 90 per cent.

per annum would be necessary to keep the sparrow population

stationary. Nodoubt many sparrows fail to mate successfully, but,

judging from the numbers of young birds, it seems highly pro-

bable that the annual produce is a least a hundred per cent., which

implies a normal death-rate, in a stationary population, of at least

50 per cent. On January 1, 1910, the number of vertebrate

animals in our Gardens was 3186, and during the year, 2354 were

added to the collection, making a total population of 5540. In the

same period there were 1554 deaths, that is to say, the death-rate

was about 28 per cent. Here, then, are three widely different

rates, an actual percentage of 1:5 in the case of human beings, an

estimated percentage of 50 in sparrows, and an observed percent-

age of 28 in the assemblage of vertebrates in the London

Zoological Gardens. It is plain that the rates are not comparable ;

two of them refer to particular species, the third is a composite

made up from many different species of mammals, birds, reptiles,

batrachians, and fishes, each with their own different rates. The

composite percentage is an arithmetical figure as meaningless as

would be the sum of so many cows, coins and cabbages. Even in

the case of a single species, however, a generalized death-rate is

complex and misleading. The human death-rate is nearly 20 per

cent. in the first year of life, falls rapidly for the first two or three

years, and then more slowly till it reaches a minimum, and then

gradually rises as advanced age is reached. ‘The tables of

“‘ expectation of life,” calculated for the Life Assurance Companies,

show that the male infant under a year old has an expectation of

4] years, that this expectation rises to a maximum of 51 in the

fourth year, and then gradually and almost without irregularity

decreases until the age of one hundred years has been attained.

It is clear that in the case of the vast majority of animals, and

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 427

notably in the case of wild animals that have been captured and

brought into captivity, we have but vague materials for the con-

struction of tables of expectation of life or for attaching any value

to death-rates. We know roughly that there is an infantile

period of low viability from the close of which through the periods

of youth, maturity and senescence, there is a gradually decreasing

expectation of life. But there are very few cases in which the

durations of these periods or the potential duration of the whole

life are known accurately, and still fewer cases in which the

position of any single individual on its cycle of life can be deter-

mined accurately by physical signs, at least in the livinganimal. At

present there is not enough knowledge on these matters to render

possible the construction of any standards against which particular

cases could be measured.

Fortunately there remains a mode of estimating the durations

of life in a collection of animals which affords some indications of

the total effect of the environment on the duration of life. In

1870 Sir Ray Lankester * (whose intellectual acumen has un-

ravelled so many tangles in biology) drew some important

distinctions in the significance of the word longevity. Strictly

speaking, longevity denotes the duration of life of an individual ;

in practice, the word connotes some idea of relatively long duration

and is used to indicate the duration of a life that has extended

to its natural limit. A further distinction is necessary.’ The

natural limit to the longevity of the individuals of a species may

be taken to mean the average age attained by the normal members

of a species living under the conditions to which they have

become adapted by nature. The specific longevity would be the

expectation of life at birth of a normal individual of the species.

It is determined partly by the constitution, but still more by the

accidents, enemies, diseases, and other external conditions to which

the members of the species are naturally subject in every stage

of their existence, and it is modified by the powers of evasion,

protection, and resistance which they have acquired. On the

other hand, it is possible to suppose that a member of a species, by

good fortune or by artificial interference, has been removed from

the hardships natural to its lot, and placed in an environment

relatively ideal ; under such conditions it would survive the

specific longevity and attain an age which Lankester called the

potential longevity. If we regard the population of Great Britain

as living in an environment to which it has been naturally

adapted, and the inhabitants of Great Britain as representing

a species, then their specific longevity, the expectation of life at

birth, is somewhere about 50 years. What the potential longevity

is we do not know, but it is certainly more than a hundred years.

The disparity is probably much greater in other cases. In the

case of the sparrow the specific longevity, the expectation of life

* On Comparative Longevity in Man and the Lower Animals: London, 1870,

428 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

of the young sparrow when it is hatched, on the most favourable

estimate cannot be more than one or two years; the potential

longevity of small Passerine birds is certainly over 20 years.

The difference between the specific longevity and the potential

longevity is a rough measure of the severity of the natural

conditions under which a species lives, and a similar mode of

comparison can be applied to assemblages of creatures living

under different kinds of unnatural conditions. The object of this

communication is in the first place to suggest a fashion in which

a system of this kind could be employed usefully in Zoological

Gardens, and in the second place to give the results of examina-

tion of a series of records regarding the duration of life of

mammals and birds in captivity, which, although they are

imperfect in many respects, are more extensive than any with

which I am acquainted. The records in question are contained

in two manuscript folios, kept in the Prosectorium of this Society

and covering the period from 1870 to 1902. In these were

entered day by day the names of the animals that had died in the

Gardens, with the dates of their arrival and of their departure.

From the point of view of estimating the effects of captivity on

duration of life, they are defective in two important respects. In

the first place, there is no record of the ages of the animals on

arrival. This of course must always be impossible in the vast

majority of cases, but two stages, the infantile and the senile,

ought to be noted and excluded, where possible. Of these, the

senile stage is usually easy to detect in the case of mammals, and

it is at least probable that senile mammals are seldom purchased

or accepted. On the other hand, senility in birds is extra-

ordinarily difficult to detect, and the presence of such individuals

will probably always confuse the record. The infantile period,

corresponding to the first four years of human life, but of course

varying greatly in its duration in different animals, is easy to

detect; and as the difficulties of rearing infants are entirely

different from the general problems of animals in captivity, infants

should be excluded from records such as those with which I am

dealing. Ina number of cases, but not in all, I have been able

to discard “infantile” entries. The second important defect in

the statistics is that they do not include any note of the condition

of the animals on arrival. Owing to the mode of capture, and still

more to the conditions of transport, a large number of wild

animals arrive at Zoological Gardens in poor condition, and die

from causes not due to their new environment. For the kind of

investigation I am now discussing (as of course for other reasons)

new arrivals ought to be kept in quarantine. The duration of the

quarantine need not be for a fixed period, but should be long

enough to make certain that there has been recovery from the

effects of capture and transport, and the date of detention in the

Gardens should begin with that of liberation from quarantine.

RELATIVE VIABILILY IN MAMMALS AND BIRDS. 429

With these comments I shall now pass to examination of the

records, which however imperfect afford a considerable body of

information *, ‘

MAMMALIA 7.

QUADRUMANA.

Average Maximum

Sen sndividuale “hy aoa

Anthropopithecus troglodytes(Chim- — 20 8 46

panzee).

Zils, CONSID eg bie ee epee eee Ree 2 61 94

Atrgarilar (Gorilla) ess: sansscsse 6: 2 30 5

Suma satyrus (Orang) ...../......... 20 DD 21

Hylobates (Gibbon), whole genus... 39 9 51

JEL CODCOD ete CAME STC Sa 9 17 51

TELAT WR eta MOE ME Tee OI 16 Ad 12

VERT LOS Gis SRC OO Oe 1 Le ORE 2 5:5 9

STEM AUICISEUES 1 Sai Renn ere ore eee 6 veil 29

VEC MCOGEIY Sha ot crear iatt a bidet te 1 7 fl

Vs RGA ei eee wo ee Dene ce 2 2°5 4

LEDGE TOPS Gee Ceo nee i 6 6

TUES OUUCCY EUS. os 08s cake vo iei ass vas il S) 9

1a bn OTH TERED TOU She be ge era 1 13 13

We know that the figures in the third column are very much

below the potential longevities of these animals, whilst the

differences between the second and third columns are equally

notable. It cannot be doubted that the conditions during the period

in question were unfavourable to anthropoid apes. In 1901, after

the period to which these figures relate, the existing house for

Anthropoid Apes was completed. The newconditions differed from

the old in providing more space, better ventilation, and protection

from improper feeding by the public. These changes have been

beneficial; in the case of the chimpanzee, of the orang and of

the gibbon, there is at present alive in the Gardens at least one

individual that has lived longer with us than the longest-liver of

the previous period. On the other hand, the average duration

has not risen notably. These animals must be regarded as of

relatively low viability, a conclusion perhaps less surprising if it

* Since this paper was written I have seen the extremely interesting “List of

Animals,” living in the Zoological Gardens at Giza, near Cairo, 2nd Edition, and

published in 1910. ‘The author, Capt. Stanley S. Flower, has made the useful

innovation of inserting opposite each species a note of the maximum ascertained

length of life of an individual specimen in the Gardens. I have added to my paper

some of Capt. Flower’s records, but as these cover a period of onlv 12 years, higher

maximums were impossible; if he were to publish average durations and state

the general conditions of captivity of the different species, the work would be

invaluable.

+ The names used in this memoir, as far as possible, are those employed in the

ninth edition of the “List of Vertebrated Animals now or lately living in the

Gardens of the Zoological Society of London”; I have used Trouessart for

determining synonyms.

430 DR. P, CHALMERS MITCHELL ON LONGEVITY AND

be remembered that the lower races of man are of lower viability

than the civilized races. None the less, existing results are so bad,

that drastic experiments would be justified. The simplest change

would be to allow them free access to the open air.

No. of Av. dur. Max. dur.

CERCOPITHECID, Indiv. inmonths. inmonths,

Semnopithecus, whole genus ......... Se OO 4:3 15

[Sie Grntd aes (Ibeuneaiiie)) Waer Wao conan ssodceeces 20 4-6 13

SNe Ga NOU UARUSISA. 5s donee podbean boadsegonns 24 3'6 15

Sq MOCHURUIS — qasuuunoocanecse. Eeadgasy savas 8 4 15

SLOUSCTTUS Geer e a) sth Receaysat beer gige sane 2 12°5 14

WS GLUPOLERL CUS Septet sack erase 1 3 3

Here again it is plain that the conditions are hostile, and that

these monkeys have a low viability. In the natural condition

they feed chiefly on grain, and green leaves and shoots, and it is

probable that their diet in captivity is not sufiiciently bulky in

proportion to its nutritive value. But these monkeys have always

been treated as requiring warmth rather than fresh air.

No. of Avy. dur. Max. dur.

Indiv. in months. in months.

Colobus, various species, chiefly

WOUETOSUS RACE, ete 10 32 13

The African Colobus and Guereza monkeys resemble the Asiatic

Semnopitheci in structure and habits. They appear equally to

suffer from the conditions of captivity.

No. of Av. dur. Max. dur.

Indiv. inmonths. in months.

Cercopithecus,* whole genus ...... 693 115 22

CPT ee ee 42 13 45

CRiUEGIEC LAGI REA UNTO il 40 40

OS MAUL OU ED: Jentnocboananaancscnhodes ug 9 44

CHRICLEATISM eee ee 17 8 30

CLARET. Cen 3 22 122

COURT AETES ERE A TEE Bae Re 1 ey es

Costas Be ee 4 8 G2

OL HUIDOC MICOS: ondascookssogon5eq0000K 45 ial 46

OES CLRUIIOM Germ a eee Tocnadtie e 6 12 Al

CRUNCH RR, EM PNR Vl 28 7 65

ON campueliy . Ste hen eet: 27 12:9 76

OU OCS UR OE id ly ER Rui Oe etek 1 44 44

OR DAU OH POSIG Marmion’ ieaadodcnodnonans 39 Mer 66

CHOUSCONTAS es eee peek EN ey RA 2 ai al

CA SEMA EO RAE See 2 3 5

CREEPS CRNA NEO MORIN od Hee 37 17-7 65

CORCTYLNE GUIS) cats eMC econ ee 2 33°5 47

* Jn this genus, I have followed as nearly as possible the synonymy in Mr. R. I.

Pocock’s revision, P. Z. 8. 1907, p. 677.

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 431

EE AIRC REE (Ga) ie ee ee etonchs.

Cercopithecus Sabeus .........0.000s 109 th 43

(= callitrichus)

C. wethiops (= griseo-viridis) ...... 24 14 60

OS ETD LLL OES Se 59 13 87

CREED) IS ese wala eas ndoia “eins sae ee «s 5 15 ot

C. pygerythrus (= lalandit) ...... 146 8°6 59

Cia ICT EOI I Se A Beane 15 13°8 34

Cin J DLLLTS ROLE AEA Eat We Ree 40 6°9 27

GP DYTTLOTOUUS Se. 5.008 Ake a gue oe 9 5b 16

Cercocebus, whole genus...........- 85 12°2 48

(Mangabey.)

Oe EUG PSE so acta nae ais Sue diic Gconelans 15 7:5 48

OPS TLULOCIED Seen rene Kaas rayne a shrs oR: 15 LOS 42

OM MLTRUIUUS aon de estas ee it 5 D

(Ooi GOLUGT EOS Mie ta e ae ors Se 15 12 38

OMGhTUSOGUSTER” | x cnes ages scene 1 12 12

ENE LEOILOSU Sie Na OM oa) 998 36 16 40

(Ors) GHG (STITT! Sake Oe eet OE Ee 2 8 10

The Cercopitheques and Mangabeys for the very large number

of individuals recorded thus show an average duration of about

one year; the best record, about ten years, must be taken as far

below the potential longevity of the group, and it is clear that the

conditions of captivity press hardly on them. Throughout the

period in question, they were treated as animals which must be

protected against cold, and they were allowed no access to the open

air.

_ No. of Ay. dur. Max. dur.

Indiv. in months. in months.

Macacus, whole genus ............ 910 14 143

(Macaques. )

DMR OSSOTCTISIS cod si: cece csaacteesais es 1 5 5

WE OANA UCU Ss 2998. i Pee seis ss os 5 12°8 22,

YVTESET/ CLO TTS OA ct Necscisclbins 2 20 26

DUEL CYTOTLOLGUS! Vege. cet et eee ea 328 16°5 130

WU TUAESTOUNIS) Wee ae Rare 3 28°6 4]

Ui le IAT TOOT IO AS Sek eRe RE |2 > RRR 7 18 rar |

IE OMVESERUNALS) cic ccricienioten see 58 20 62

JL TOAUD OS Salis saa eae te oe en eee 3 10 16

JUS VOICI Sener BOOS aor AOR 21 10 atl

DER CS UUSIM NG | ses ic otieatede ae cic cians’ 22, 16:6 143

BL (tO HOCTOSD ee 3 5:3 vy

WSO HIEC nae» anne =) Seren see 16 30 119

VEE SUES Mee oh Sos Sisilred Pa < tested Gee 203 13 75

WES SHE CLOSILS Mee ae his sae HOMRs chica 4 4] 104

MEANT US: SARS se cliccicoetencceelsee’ a 18 fia

LMT ROUCOCKAS 5 SHS RAOO RBA O OD ona Bees 2 16°5 19

VES Gf ESCONS FA stk. onomancee anew 2 8 15

VSS OCT.COLUS) aie see ee eee ) 1] 28

MBO VAUUSS:. CUI SO a ee 24 12-3 38

432 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

The large number of Macaque monkeys recorded shows the

same low average duration, and the same marked difference between

the average and the maximum duration, although the latter is

certainly below the potential longevity. A Common Macaque

occasionally deposited temporarily at the Gardens is stated by

its owner to be at least 22 years old. It is of significance that

the highest average relates to Japanese apes, examples of which,

unlike all the others, were housed in a cage exposed to the open

air.

No, of Av. dur. Max. dur.

Indiv. in months. im months.

Cynopithecus WAger .........0.0--0 00 9 10 Al

(Celebes Black Ape.)

All Balboomst ip bacncccctsnm toateenca: 137 12-2 113

Theropithecus gelada@ ............--- 2 9-5 10

Cynocephalus anubis ............... 12 12°6 51

CECA DOI: ON I AVE IOS: 19 10 Its}

OCRAOG ETE PR Ree 1 4 4

Oh JODO UF OCS 000000008800 seneoenG0% 22 12 29

CRUE CTIISIMES Ae PON OR INE OTRO 1 1 1

Oleicophatseins Ae tet. alee 6 16 26

OG MOMOOD sosbus seo seaccbseads5s600 ns 7 15 25

(OSM IGIUCH IO), “seaadoreneeonooeecs a 1 7 7

ON DAP COW 6A Rh I eR BB 2 3 6

OR DORCU RUSH: <A: Lee eee 43 12 50

ORES) 0) COMO See oot Gon dace me Coen cer 21 15 A]

Baboons, on the average, show even a lower duration and a

greater difference between the average and the potential longevi-

ties than Macaques. Since the period in question, a number of

baboons have been exposed to the air and the average duration is

notably higher.

No. of Av. dur. Max. dur.

CrBipa. Indiv. in months. in months,

Chrysothniaisciured .aane...s0., 44 9 108

(Squirrel Monkey.)

Cebus, whole genus ............,.. 226 10 50

(Capuchins. )

CR GUD O SHEN ca ereva Renee iar 33 8°6 20

ORG DOUG NWN chen ae 10 11-8 Dit

CSD EUEOY (SRN oa. Fe 2 11 16

(OF (QTV DUICOLLCIS (WARD OP Roe une. Sheoasene ee 39 75 20

CRG QUMCUUWS ER cies tke atom aateeet 58 11°8 50

CRUUGUCSCCTISOR Jini ccinnn Re oe 6 8°D 12

Ol VOT OTEDUGIIS sana scli 54> sdhaocoocds 46 8 17

CRUNCHES. ee Re ee 9 We 39

CRIN ORUCHUSMM ore eee eee Ee 21 14:3 AQ

CiSfoellenos nist secs accent erie tetet 1 15 15

OROUIRIODIUIP. udsosodoooos ooavesadeass 1 31 31

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 433

No. of Av. dur. Max. dur.

Indiv. inmonths. inmonths.

Ateles, whole genus. ............... 107 5°8 27

(Spider-monkeys.)

EAGLE Pp Cesta et ot ORT ne) Aa eee eee 54 4:5 27

LIANG COIPENOU Da tes», tl sancs cin eiarine, Bongo d 10 (es) 24

WAN GUISE SCO MUS tapes des slats « scduin/atsieh otogalt ope 1 2 2

Beg OPIOUSE A ca senjek Rend de okie oi, ] 9 9

PART UGITO CIT iad iat sd aelossisratge cfosterf: 31 8 20

TTT DESGUIS PE ay, Ob RO ean Cee Oe EE 4 5:2 10

AAT UP UCIUERIS sv hecoptnte strain’ ed baie lem 2, 3 4

AS CTLOG ADU Stefi jerdoishinrs aeinacyatt dese 3 46 11

PA MUCLICT OSU Siig 2 ais artousx 46, sites Ayapeiayen' 1 3 3

Lagothrix hwmboldti ............... 14 5:2 16

Nyctipithecus, whole genus ...... 27 13 AD

(Douroucoulis.)

INE AIC cen ookb Ope enact ce SURE eRe 14 13 45

ING GOGIYOIS. ented wpeoneoceeds: > eeE Nees ] 1 i

NEMERCUUNGGLUUS) Sa ete iyeecr eae a ars. 2 9) 5

ING SUOCI Er UIS: Pere «sete hea: eo << 10 15) & 98

Callithrix, whole genus ............ 5 4 8

(Teetees. )

Pithecia, whole genus ...........-. 16 Zia i

Ouacaria, whole genus ............ 7 3°5 11

Mycetes, whole genus. ............ 9 3°6 13

The Cebide show the same marked disparity between the

average duration and the maximum duration, although again the

latter must be considerably less than the potential longevity. It

is interesting to notice that the Capuchins and Squirrel Monkeys

display a higher viability than the others.

HAPALIDA. No. of Av. dur. Max. dur.

(Marmosets. ) Indiv. in months. in months.

Hapale, whole genus ............... 232 3°8 16

JER OUKISOLANIGDS. Socosevocee eee ee 1 i if

ELS OCCN IES, WNP Nephi. 0 AAR Rese otek 134 4-2 16

Yale IGG OUIRLL, 85 Core oe net 6 a cesdecs. 1 6 6

ETD CRC CECE tN aa er tlle hc 94 34 16

TTY GUE Us: ANE chen. Bas oP ecvartaine 2 2°5 5

Midas, whole genus .....2..2..+++. 103 6 110

MR GhpySOMClasy hn soasttem tek dekeke 2 6°5 1]

MEROCO POU Bit ets Verh PAE: otha a 2°4. 4

WM LOCUGL CUS ek ROR oor nan ated atyece 1 4 4

AM SUSE AC ieee Pods 4 sc Belge! ele. 1 5) 5

MiSs UU PUis cae sack erica GO as Ss,s 25 4 14

Mes nOSaliGk Vinge S244 septsas ake 39 9 110

DE RU UIICH OLS aoe done enone? Rao nee 11 45 16

Wie aiasanhit Shp eM ape cadte. Sie eet: 17 4:6 12

Amongst the numerous Marmosets dealt with in the record, a

A434 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

single case of relatively high longevity, that of a Lion Marmoset

which lived in the Gardens for over 9 years, has given those of the

genus Midas an average duration of nearly double that of the

representatives of Hapale. Hven this fortunate instance, however,

does not disguise the low viability of Marmosets under the

conditions of their captivity. As these animals spend a large

part of their time in their sleeping boxes, unwise feeding by

visitors cannot account for much. On the other hand, they have

been treated as animals that required special protection from cold

and fresh air.

Miss C. Morey informs me that she has kept a marmoset for

16 years; it was allowed great freedom and had constant access

to fresh air.

LEMURES. No. of Av. dur. Max. dur.

LEMuURIDA. Indiy. in months. in months.

Lemur, whole genus ..............: 155 25°3 134

DONO ROO Wan seaoeecalon ses Sear ae: 16 43°6 113

She, UOWROTUS sesogocsscancesosoesneec 1 36 36

LE OP CHODUC TIS atoies ony Oo NOH ve oe 22 25 98

HES COULG dia NAR Hela ON tu inl 33 i 38

Loy COPOMAHO US fo ncisedadsoededsosbedaeeno: 10 G5 5D

Lin UDC POMS Wwabcondosardsesdecs dass 1 AD 45

UTR TVUCHC Orin tee pe tN OP Ihe 46 134

Jb GHEIOTAOS Sogo 0ecesenontesneo vee: 1 2 2

WES ROR OR oe eticeh bee aasten eek Ree 7 39 122

TRG CT Tec AEs PRONE AE = AOR. 3 63 107

JL; GUTEPPUTOLES! conooc den scosAnh rab odooe 2 30°5 43

TREAUOPUF EONS 1) Tan MEN, SO ED 3 23 46

TOU TONS EEN Aa te EROS, E 13 27 113

LLG UJUDOS crete ES ee 3 22 29

SU GTV US RE Levee sei heen eer Nene 10 20°9 77

Jb FETAL OMDORSITEG neon goes 2066068656 13 29 72

Llapalemurrstnus .. 5.0.2.8 2 DD 8

TEL SGRUSE US BER Bote RRS 6 22 58

Chirogaleus coquereli ............... 8 82 184

CGR AR RN Vick WE 2 ih, 2 70 138

WMicnocebusismith? sa ene eee 14 20 84

Galago, whole genus ............... 61 22 86

GST ei 4 Ninn el ance at ae 3 9 15

EO NCRUSSCUUAALA ener ee 7 16 34

G NG ONTICEL I: ARM 15 ine Nar Slee 20 175 A3

GAT CIVOU CA gk aati e eR ee Ee 25 26°7 78

GE ATO NECUTIM 2S). Se kee REP ERA 6 32 86

Nyecticebus javamicus ............... 1 21 21

J, GORCMOTROUUIS. Woonssecsadnoos0s00u 29 13 AQ

TGTISKG OCU IS wane ate ee 12 374 7

TET ROONEIOS JULIO.“ Gandnooconsnonsces 18 9°3 22

Chiromys madagascariensis ...... 3 36'3 107

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 435

With the exception of the Loris and the Potto, the Lemurs

show a longer average duration and a higher maximum duration

than in the case of Monkeys. There still remains, however, a

very marked discrepancy between the average duration and the

maximum duration, although the latter must be regarded as much

below the probable potential longevity. On the whole, lemurs

have been kept under conditions not very different from those of

the monkeys. It is probably in their favour that as they are

more nocturnal than monkeys, they have escaped to a greater

extent erroneous feeding by the public, whilst as they are on the

whole more strictly arboreal than monkeys, the relative absence

of light and the more equable temperature in the interior of

warmed houses have not been so damaging to them. On the

other hand, they are, on the average, smaller than monkeys, and

as within the same group smaller animals have usually shorter

lives than larger animals, a longer average duration and a

higher maximum duration as compared with monkeys, was not to

be expected.

Summary of Quadrumana.

It is well known that the higher races of men have a higher

viability than the lower races; that is to say, they display a

greater power of resistance to adverse conditions, to changed

conditions, and to diseases (although not necessarily to any

particular disease). It is of interest to find that a still lower

viability is exhibited by apes and monkeys which as a group

have an amazingly poor power of resistance to the adverse con-

ditions of captivity. It is even more interesting to note that

there is a marked gap between monkeys and lemurs in this respect.

The fact that the latter on an average live four or five times

as long in captivity as monkeys, shows a marked difference in their

constitutions. I may recall a curious by-product of an investi-

gation into the feeding of snakes made by my friend and collea gue

Mr. R. 1. Pocock and myseif (P. Z. 8. 1907, p. 785), when we found

that lemurs differed from monkeys in having no fear of snakes.

Fundamental differences between groups are revealed not only by

anatomical characters, and in view-of the present tendency to lay

increasing stress on the anatomical links between lemurs and other

primates, I think their physiological differences are worth some

attention. There is, however, another interpretation of the

higher viability of lemurs in captivity. The close affinity of man

and monkeys may render the latter more subject to human diseases

and therefore more lable to suffer from the vicinity of man. Such

a question can be answered only by long records of the causes of

mortality based on adequate diagnosis. The laborious work of

my friend and colleague Mr. H. G. Plimmer, unfortunately does

not extend back to the period with which my present inquiry deals.

Even if it be accepted that apes, monkeys, and lemurs are

creatures of inherently low powers of resistance, the striking

difference between the average durations and the maximum

436 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

durations makes it clear that they have been subjected to adverse

conditions. Throughout the period to which the figures relate,

and in a majority of cases since, the conditions of their housing

have been determined by considerations of temperature rather

than of fresh air. The conclusion is inevitable that these con-

ditions are adverse. This is not the place to discuss the

construction of houses for animals, but it may be well to say

that there is no reason to suppose that the other extreme, com-

plete indifference to the provision of warmth; would be much

better, although it could not be much worse.

Capt. Stanley Flower’s records show no figures of maximum

duration greater than those recorded here.

CARNIVORA.

FELIDE. No. of Ay. dur. Max. dur.

Indiv. in months. in months.

Gh ao (LAGI) ened. dobeosvasdawnd aes a5) A5-4 204.

HH SGHOMIS (ISON) he eka ee eee 30 44-2 145

Jil, yoapainis (QUE PARC) 52.420 sc 255202040: 52 39°9 164

FF, uncia (Snow-leopard) ..... rae 2 20°5 AO

Jd sO 0o( (wig) chgeee th ce ecole 8 57 169

JES GGG (Loe LyPIOwS)) eodevasolsess os888 3 DD 122

fF, canadensis (Canadian Lynx) ... 2 13 139

it amare (Ceneeall)) 5 ddosabiesieensoe- 9 63 131

HE coneotor, ( uma) nt). Arcee eopivints 18 A\-2 124

JTS O10GT (VESTA) Poecodssosaodacsosdseo 8 33°5 123

I have omitted from these figures a certain number of entries

of under one month and which, in most cases, related to very young

animals or new-born cubs. ‘The highest record in the period,

17 years, in the case of a Lion, is certainly very much under the

potential longevity of the group, for lions of between 30 and 40

years old have been known. The discrepancy between the

average duration and the maximum duration is still great, but

very much less serious than in the case of apes and monkeys, and

considerably less than in the case of lemurs. Capt. Stanley Flower

gives no higher records.

No. of Ay. dur. Max. dur.

Indiv. in months. in months.

F. nebulosa (Clouded Tiger)......... 1 62 62

Hasenvaln (Servall) Ms. Aes 4 secceete oct 22 21-7 98

F. servalina (Servaline Cat) ......... 9) 9 AD5

F. temmincki (Golden Cat) ......... 4 24 65

I, viverrina (Viverrine Cat)......... 4 51 112

F. bengalensis (Bengalese Cat)...... 22 16 77

fF. planiceps (Rusty Tiger Cat) .... 2 9) 5)

F. rubiginosa (Rusty Spotted Cat). 2 15, 2

F, chrysothrix (Red Tiger Cat)...... 1 1 1

Liyeatis(NValdg ait) aes. ecko. 31 15 110

fF. chaus (Jungle Cat) ............... 15 AT 118

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 437

No. of Ay. dur, Max. dur.

Indiv. inmonths. in months.

felis caffra (Kather Cat) ............ 3 1 2

F. maniculata (Fettered Cat) ...... 2 17:5 35

F. yaguarundi (Yaguarundi Cat)... 5 1 3

PERCU ( WAv CARO Alby) eines acs farsal seie\e' 3 10 19 109

FP. pardalis (Ocelot) .................. 56 U7 152

F. tigrina (Tigrine Wat, ise. 4 3 i,

F, geoffrott (Geoftroy’s Cat) ......... 10 17°8 103

F. passerum (Pampas Cat) ......... 2 8 10

The records of average and maximum duration of these smaller

Cats are very irregular, and in many cases the numbers are too

small for the result to be more than accidental. We do not know

what is the potential longevity of the smaller cats; it may be

expected to be less than that of large cats, and has been estimated

at something over twenty years. It is clear that the difference

between average duration and potential duration has been much

greater in the case of the small cats than in that of the large

eats. The domestic cat is notoriously a hardy animal, and I

should hesitate to infer a lower viability for small cats than for

large cats. In the first place, throughout the period under dis-

cussion, the majority of the large cats were kept in the Lion House,

a building i in which the ventilation and bulk of air per inhabitant

were fairly g good, whilst, although there was not free communi-

cation with “ae outer air, there were outdoor ¢ cages, to which on

occasion the animals had access. The smaller cats, on the other

hand, were kept in a smaller and darker house, <ttlh much less

ventilation, more heating, and no access to the open air. In the

second place, there is an Fmportant. natural difference with regard

to feeding, which has not been allowed for systematically. The

larger cats are accustomed to make an occasional kill, and to

return to a carcase even after it is putrid. The smaller cats, on

the other hand, devour their smaller prey as soon as they have

killed it, and unless under the pressure of serious hunger, take

nothing except quite fresh food. In accordance with their natural

habits, the larger cats are little injured by food which is not quite

fresh, whilst the smaller forms are highly susceptible to intestinal

ailments arising from the quality of their food. Iam convinced

that the necessity for extreme care in the quality of food given to

the smaller cats has not been sufficiently realized.

I do not think there can be any doubt, however, that both with

smaller and larger Felidz, the necessity of access to the open air,

at the choice of the animals, has not been appreciated, and that

the low viability under menagerie conditions is due to this neglect.

Most of them are partly nocturnal, many of them cover a wide

climatic range, and as in many, if not all furred animals, the

thickness and length of their coat increase rapidly with exposure

to low temperatures. No doubt provision for basking in the

sun, or for warming themselves in artificially-heated retreats, is

Proc. Zoot, Soc.—1911, No. XXIX. 29

438 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

necessary in most cases, but even more necessary is constant

access to the open air, by night and by day, at the free choice of

the animals.

No. of Ay. dur. Max. dur.

PrLips (cont.). Indiv. in months. in months.

Cynelurus jubatus (Cheetah) ......... 17 30 78

Cryptoprocta ferox (Cryptoprocta) ... 2 A-5 5

Capt. Stanley Flower records a duration of over 9 years for

a Cheetah, the animal being still alive.

VIVERRID. ene hued. inane

Viverra, whole genus .........-...-......2: 19 65 181

Ve civetta ( Atricam’ Civet)) 2.0.42. 2255: 14 66 181

V. tangalunga (Sumatran Civet) ...... 4 40 81

V. zibetha (Large Inchan Civet) ...... 1 143 143

Viverricula, whole genus ............... 12 33°D 93

V. malaccensis (Indian Civet) ....... a SliD 34°5 93

V. schlegeli (Schlegel’s Civet) ............ 1 23 23

We have no information as to the potential longevity of Civets,

but the record of 15 years is no doubt considerably too short. On

the other hand, their average duration of life has been good; it

is interesting to notice that, although these are at least as typically

tropical animals as the majority of the Felide, they have been

for the most part kept out of doors in unwarmed shelters.

Genetta, whole genus ........40..--.2++-+- 46 30 148

Gajelinal(MelameyGenet) ieee. .ceee.- ee 2 16 16

G. pardina (Pardine Genet) ............ 4 AT-5 148

G. senegalensis (Pale Genet) ............ 4 8 17

G. tigrina (Blotched Genet) ............ 22 35 122

G. vulgaris (Common Genet) ......... 14 27 101

Paradoxurus, whole genus ............ 58 35 185

P. aureus (Golden Paradoxure) ...... 3

P. gray (Gray’s Paradoxure) ......... 3 5D 99

P. niger (P. prehensilis, P. pallasi, 35

P. hermaphroditus) (Common

Paradoxure).

P. larvatus (Masked Paradoxure)...... 8 64 185

P. leucomystax(White-whiskered Para- 4 D3 166

doxure).

P.philippensis(Philippine Paradoxure) 5 28 102

Arctogale leucotis (Three-striped Para- 8 40, 132

doxure).

Astnioingate | tenssP se wan ase 1 D 5

Hemigalea hardwickii (Hardwicke’s 2 55 i

Hemigale).

Aretictis binturong (Binturong) ...... 8 39 82

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 439

No. of Ay. dur. Max. dur.

Indiv. in months. in months.

Nandinia binotata (Two-spotted Para- 16 32 85

doxure).

Galidia elegans (Hlegant Galidia) ...... 2 Ad 9

Herpestes, whole genus .................. 152 20°8 105

H. albicauda (White-tailed Ichneumon) 3 Biles CE

H. fulvescens (Macearthy’s 3 aes 15 20

Hf. galera (Marsh i 9 31 57

H. gracilis (Slender a Dee 5 17-4 35

H. griseus (Grey ‘, a) 14 53

HH. ichnewmon (Kgyptian i els AQ) gu

H. awuro-punctatus (Spotted 33 ), 30 18 57

H. pulverulentus (Dusty FF Nias 47 105

HE, smithi (Ruddy i red. 22 35

Cynictis penicillata (Levaillant’s 18 32 145

Cynictis).

Crossarchus obscurus (Kusimanse) 5 15°6 AQ

C. fasciatus (Banded Ichneumon)...... 24 32°8 92

Suricata tetradactyla (Suricate)......... AD 12°6 58

The Genets, Paradoxures, Ichneumons, and Suricates are small

carnivores varying in their diet from purely carnivorous (in-

cluding small mammals, birds, reptiles, eggs, insects) to a mixed

and partly frugivorous or vegetarian type. What their potential

longevity may be is quite unknown, but the records of maximum

duration going to over fifteen years are unexpectedly high for these

small animals. The average duration varies very much, but is

lower than in the case of the civets. During the period covered

by the records, the genets, paradoxures, etc., unlike the civets, have

been kept in most cases in a warmed house, without access to the

open air.

Capt. Stanley Flower has no higher records.

PROTELID#. No. of Ay. dur. Max. dur.

Indiv. inmonths. in months.

Proteles cristatus (Aard-Wolf) ......... 6 53 155

We have no information as to the potential longevity of this

aberrant animal, but it appears to have a high viability.

Hy ZNIDE:

Hyena, WHOLE BENUS s......25. 060.50 43 29°5 160

H. brunnea (Brown Hyena) ......... 5 30-2 56

H. crocuta (Spotted ,, )......... 11 58:7 160

EL SirigtGe g(SbrIPeds cr |) osnsc2a5. 27 17°6 74

Nothing is known as to the potential longevity of hyenas, and

the record of 13 years in the Gardens may possibly be a fair age

for these animals. They have been kept at the Gardens in cages

under cover, exposed on one side to the open air and provided

29*

440 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

with hot-water pipes. They have had little room for exercise.

The lower viability of the striped species is difficult to explain.

Capt. Stanley Flower quotes a Spotted Hyzna and a Striped

Hyena, each with a duration of over 10 years, and still alive.

CANIDA. No.of Av.dur. Max. dur.

Indiv. inmonths. in months,

Canis lupus (Common Wolf) ......... 18 29 72

C'. hodophylax (Japanese ,, ).........--. 1 90 90

CO, woalloes. (Wan eben 44 \socconeaseoe 5 42 149

Ol, hetimaos (ETN. 4) neosnseanses 4 59 115

C. antarciicus(Antarctie,, )...-.---.--- 2 A415 64

CO. niger (Black Sy laceenorenene 2 75 90

AU WV OIVOSWA.. take Mata d Saja etc: 32 AQ 149

C. anthus (North-African Jackal)...... 36 Bes) 82

C. aureus (Common Sel) eee 16 22-4 122

C. lateralis (Side-striped “seh aetess 8 24 56

C. mesomelas (Black-backed ,, _ )...... 39 19) 164

PACK ENS Iaes aces. ce beers niise eee ace ee 99 18 164

Or dG (DINE) 9360026 eeaa se acnissace i 43 145

C’. (Cuon) primevus (Indian Wild Dog) 6 35 a)

C'.(Cuon) rutilans (Malay Wild Dog). 4 22 28

C. jubatus (“‘ Maned Wolf”)............ 2 15:5 23

(Ol, Ceaearag (ame DOS) so505s00ne0008 20. 1g 39 103

C’. cancrivorus (Crab-eating Dog) ...... 19 15 60

C. magellanicus (Magellanic Dog)...... 4 32 100

C. fulvicaudus (Red-tailed Dog) ...... 1 53 53

C. fulvipes (Red-footed Dog)............ 1 16 16

C'. (Nyctereutes) procyonoides (Raccoon 13 26 69

Dog

C’. microtis (Small-eared Dog) ....... : 1 4 4A

Mlitfoxclike Doest pcs cutee vies 58 Dil 103

C. argentatus (Silver Fox) ............--. 1 48 48

C. bengalensis (Bengal Fox) ............ 12 21 68

O's CLOUD (UEIMNEG INO) > sasscocecooseseses 10 249 110

C. chama (Silver-backed Fox) ......... 15 19 62

C’. famelicus (Syrian Fennec) ......... 5 14:4 32

Of, jroo (L880) INOS) Aascscaosonoseesdocer 21 30 S77

Ol, KagajsuIs (CACMG 180s) }ocvode0 sob0des8o 24 488 132

C. leucopus (Indian Desert-Fox) ...... 6 50 91

C. niloticus (Egyptian Fox) ............ 9 32°5 CO

C. pallidus (Pale Fennec Fox) ......... 5) 15 60

Of gn tioaa | IRING INNS) p pook cosaddaoseoacobacee 5 5 34 71

C. virgimanus (Virginian Fox)......... 1] 12°6 30

C, wailleas (Clomimnein INOE)) « coobbesassnse0- 4] 14:6 87

All Foxes and Fennec Foxes......... 165 26°7 132

Lycaon pictus (Cape Hunting-Dog)... 6 19°3 D6

Icticyorn venaticus (Bush-dog) ......... 1 2°5 +

Otocyon megalotis (Long-eared Fox)... 4 A-5 8

The potential longevity of Canine animals is probably not much

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 44]

more than thirty years. Ray Lankester * has recorded a domestic

dog of 34 years; the oldest Metchnikoff was able to procure was

22 years and was very decrepid. The maximum record in this

list—a Jackal which lived in the Gardens for nearly fourteen years,

but the age of which on arrival is unrecorded,—is, therefore, not

at all unexpected. The viability varies roughly with size, the

wolves having a better average than jackals, small wild dogs

and foxes. Within the period recorded, so far as I can ascertain,

the wolves have been freely exposed to the open air, whilst many,

although not all, of the smaller animals have been provided with

artificial heat.

Capt. Stanley Flower has no higher records.

No. of Ay. dur. Max. dur.

MUSTELIDS. Indiv. immonths. in months.

Mustela, whole genus..........0.c0ece000 38 29 123

M. erminea (Common Stoat)............ 7 21 39

MM, fone (Beech-Marten) ...........--.: 5 48-4 84

M. martes (Pine-Marten) ............... tf 27 107

M. pennanti (Canadian Marten) ...... 1 123 123

M. putorius (Pole-cat) ..........0.0.065 11 10:6 26

MS nulganis’ (NWexsel) 22.0) e222 see aee 7 38°7 94

Gulo luscus (Glutton) ...............-200+- 6 A2 64

Galictis barbara (Tayra) ...........2.2..+- 13 30 95

Cavitiata(Grison)! ty, eee eee 12 Al 117

Ictonyx frenata (Banded Zorilla) ...... 1 2 2

I. lybiea (Libyan Zorilla) ............... 8 8:5 A]

Ut sonllan(Cape-Zorilla\) 298 oe 18 15 65

Helictis subaurantiaca (Orange-bellied 1 48 48

Helictis).

Mellivora capensis (Cape Ratel)......... 1 208 208

M. indica (Indian Ratel) ............... 1 152 152

Tauxidea americana (American Badger) 2 130 145

Meles ankuwma (Sand Badger) ......... 3 79 152

M.leptorhynchus (Sharp-nosed Badger) 1 9 9

M. taxus (Common Badger) ............ 26 32 137

PMIMIBENOUGRES gS See eerodhe ot pote ceco epeecee. 32 41-5 152

Mephitis mephitica (Canadian Skunk). 5 25 50

Conepatus mapurito (Chilian Skunk). 5 21 AQ

Lutra leptonyx (Small-clawed Otter)... 2 ih 10

iar (indians Otter) te ses. eecen ceo: 3 32 52

L. vulgaris (Common Otter) ............ 36 AO 87

The average and maximum durations of the various Musteline

carnivores vary irregularly. The longest duration, 17 years for

a Cape Ratel, is surpassed by an Indian Ratel which has been for

20 years in the Gardens and is still alive. It is clear that the

potential longevity of the Mustelines is high, if their relatively

* Quoted by E. Metchnikoff, ‘The Prolongation of Life,’ English Translation.

Heinemann, London, 1907.

449, DR. P. CHALMERS MITCHELL ON LONGEVITY AND

small size be considered. The apparent low viability of many of

these animals may be attributed partly to their fierce and active

disposition, from which it comes about that captured animals are

frequently young or feeble, and that after capture there is heavy

mortality from fighting and injury. But there is a marked effect

of the mode of housing; the ratels and badgers have been kept in

open-air enclosures, whilst a majority of the other forms have been

housed in small cages in dark, warmed and_badly-ventilated

houses.

Capt. Stanley Flower has no higher records.

Vf 7 ~ . .

Procyontp#. Shs ucinrorireeatisance

Procyon cancrivorus (Crab-eating 10 68 190

Raccoon).

12, DUP (UEKEEOOM)) goscnnasancossccoacer 24 50 165

Nasua narica (White-nosed Coati). 35 14 46

NV. rufa (Ring-tailed Coati) ......... 63 12°5 62

Cercoleptes caudivolvulus (Kinkajou) 30 30 75

Bassaris astuta (Cunning Bassaris). 5 10°6 29

Bassaricyon allent (Allen’s Bas- 1 66 66

saricyon).

The Procyonide are another group regarding the potential

longevity of which we have no information. The record of nearly

16 years for a Raccoon shows that the group in this respect

resembles the larger mustelines. It is noteworthy that the

average longevity of the Raccoons, which have been kept out of

doors, is very much higher than that of the Coatis, which have

been kept indoors.

No. of Avy. dur, Max. dur.

ASLURIDE. Indiv. inmonths. in months.

Ailurus fulgens (Panda)...............++ 2 32 64

UrsID&.

Unstistewino ley SemuSeeensseeentee ween (5) 68°9 400

U. americanus (Black Bear) ............ 12 (i 205

U. arctos (Browns 2. seas iaee een enn we 20 23°6 133

US, loirmalbalins (Curly 5.) soosevsscces 1 164 164

U. isabellinus(Isabelline ,, )............ 4 84 153

U. japonicus (Japanese ,, ).isciictis:. 2 109°5 123

U. malayanus (Malay ,, )....5....00. 4) 91 247

O, anes (Wollee 5) onvoacaeaced 8 102 400

OU ornatus (Spectacled jy) 1.20. 2:4.- 2 96 163

U. piscator(Hairy-eared ,, )............ vA cull 309 309

U. syriacus (Syrian sale aa gaaba 8 107°6 259

U. tubetanus(Himalayan’,, )............ 12 3 179

Melursus ursinus (Sloth Bear) ......... 9 Al 109

The duration of over 33 years attained by a Polar Bear may

RELAYLIVE VIABILITY IN MAMMALS AND BIRDS. 443

possibly approach the potential longevity of bears, for the con-

ditions provided, affording exercise, constant access to fresh air

and to shelter, and regular food, must be more natural than in

the case of most animals in captivity. The average duration

of life of the bears is probably low on account of the number of

cubs which are included and which had received unwise treatment

as pets. Nevertheless, the average longevity is higher than in

the case of any other large carnivora, and it is interesting to

notice that during the period dealt with these animals, although

not given good room for exercise, were freely exposed to the open

air, and at the same time provided with some artificial heat.

Capt. Stanley Flower’s records are much lower.

No. of Ay. dur. Max. dur.

OTARIID#. Indiv. inmonths. in months.

Otaria, whole genus’ 2 ....2.cc.00..0c00e ee 6 108 210

O. californiana (Californian Sea-lion). — 1 51 51

O. hookeri (Hooker’s ti Gy cam ft 18 34

O. jubata (Patagonian ,, )2 2 199 210

O. pusilla (Cape Sea-lion) ............... 1 151 151

Halichorus gryphus (Grey Seal)......... ihe 17 73

Phoca vitulina (Common Seal) ......... 25 20 172

Monachus albiventer (Mediterranean 2 17 34

Seal).

Cystophora cristata ( meee 7 4:5 26

Seal).

Sea-lions and Seals evidently at least equal in potential longevity

other members of the order Carnivora of large size. The relatively

low average duration is due to the very heavy mortality of young

and newly caught animals. The greatest number of deaths are

of animals that have lived only a few months in the Gardens; if

they survive that period, and only a small proportion do survive

it, their duration of life is very good. They have free access to

epen air, and no artificial heat.

Summary of Carnivora.

Carnivora are generally regarded as animals with relatively long

lives and high viability, and the records given above support such

aview. It is curious, moreover, how the maximum durations

correspond roughly with the sizes of the animals. Bears reached

33 years, Lions and Tigers 17, Sea-lions 17, the smaller Cats 13,

Viverrids 15, Hyznas 13, Jackals 13, Badgers 12, smaller Dogs

and Foxes 10, Mustelines 10. On the other hand, comparison of

average durations with maximum durations shows in the clearest

way that those animals have lived best which have had the readiest

access to fresh air, that, in fact, darkness and the bad air of

heated houses are much greater dangers to life than cold.

444 Dk. P. CHALMERS MITCHELL ON LONGEVITY AND

INSECTIVORA.

No.of Av.dur. Max. dur.

SORECID. Tae in Hee in Sata

Crocidura cerulescens (Bluish Shrew). 1 1 1

ERINACEIDA.

Hrinaceus, whole genus ............------ 53 10 54

E. albiventris (White-bellied Hedgehog) 1 1 1

FE. algirus (Algerian Hedgehog) ...... 3 28 39

HE. auritus (Long-eared ,, ) ...... 8 12 24

E. collaris (Collared et ae ete tee 6 1) 54

Hi. europeus(Kuropean ,, ) ...... 33 7 20

E. frontalis (Cape Saat g) Weeeee il 1 1

H. micropus (Small-footed ,, +) ...... 1 1 1

TUPAUIDS.

Tupaia peguana (Peguan Tree-Shrew) 1 22 22

iE canvad (hanaeAltenpania) eee sapere il 28 28

CENTETID A.

Centetes ecaudatus (Tenrec) ............ Tl 10°5 24

I do not know of any information as to the potential longevity

of Insectivora, and the few records which I have found would

appear to show that both the viability and the longevity of these

animals under the conditions of captivity are bad. They have

invariably been kept indoors, in small cages in houses heated i

winter.

Capt. Stanley Flower’s records are all low, the maximum being

34 months for a Long-eared Hedgehog.

CHIROPTERA.

No. of Av. dur. Max. dur.

PTEROPODIDA. Indiv. inmonths. in months.

Pteropus, whole genus .............+. 42 30 205

P. formosus (Formosan Fruit-bat). 3 62 94

P. gouldi (Gouldian See yok pel 58 58

P.medius (Indian et 0) 27 205

P. poliocephalus(Australian ,, ). 6 22 83

P. pselaphon (Bonin ee fa iene 26 26

P. scapulatus (Scapulated ,, ). 1 ff 7

Cynonycteris collaris (Collared 13 18 108

Fruit-Bat).

Cynopterus marginatus(White-eared 4 24 30

Fruit-Bat). °

Metchnikoff has already recorded from the London Zoological

Gardens the case of maximum duration amounting to 17 years

in a Fruit-bat, and has pointed out that the probable maximum

longevity of such batsis high. The records show that the viability

under existing conditions is not good, as wherever there is a

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 445

considerable number of specimens, the average duration is much

below the maximum duration. There is no difficulty about feeding

these animals, but they have always been treated as creatures

requiring protection from cold and indifferent to fresh air.

Capt. Stanley Flower records a Collared Fruit-bat of over

11 years’ duration and still alive.

VESPERTILIONIDE. hae a cae cane

Vespertilio noctula (Great Bat) ... 10 1:9 5

V. natterert (Natterer’s Bat) ...... 12 1 1

Plecotus auritus (Long-eared Bat). 11 09 1

Bats are popularly supposed to be long-lived animals, but these

records show only the complete failure to keep them in captivity.

The chief difficulty has been to induce them to feed.

RODENTIA.

No. of Ay. dur. Max. dur.

ScruRiIDsA. Indiv. in months. in months.

NCUURUS: WihOlesCeNUS § san-ceekeeee ee te 273 15:5 181

S. cestivans (Tropical Squirrel)... 2 22:5 Al

S. arizonensis (Arizona Dia) ie Se 39 39

S.atrodorsalis (Dark-backed ,, )... 1 10 10

S. bicolor (Jelerang SA idee et 10°5 23

S. caniceps (Hoary-headed,, )... 1 114 114

S. castaneiventris (Chestnut - bellied 2 71 107

Squirrel).

S. cinereus (Grey Squirrel) ......... 24. A8°7 181

S. ephippium (Saddled ,, )......... 1 64 64

S. erythropus(White-banded Squirrel) 2 13°5 15

S. griseus (Western Grey jie ewes 25 40,

S. griseo-flavus (Guatemala wattle: th 34 34

S. hudsonius (Hudson-Bay DB Pee) 40°5 99

S. hypopyrrhus (Dorsal By 2 344 89

4+ dorsalis.

S. ludovicianus (Yellow-footed ,, ) 2 29°5 56

S. macrurus (Grizzled Hill go) 2 WGP) 19

S. madagascariensis (Madagascar ,, ) 1 4 4

S. maximus (Malabar panne eo) 21-5 75

= indicus.

S. nigro-vittatus (Malayan po eel 34:5 68

S. palinarum (Palm 56h) uD 3-2 14

S. plantani (Plantain ale 22, 53

S. prevostt (Prevost’s Fi aduise) es) 34

S. pusillus (Dwarf bey 2 8:5 12

S. pygerythrus (Blanford’s i) a 2 2

S. stramineus (Fraser’s ey ae 8 8

S. syriacus (Syrian Pe ilvice 59 69

S. tennents (Tennent’s eat) th) D 25°4 42

S. tristriatus (Three-striped ,, ) 4 3:5 5

446 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Avy. dur. Max. dur.

Indiv. inmonths. in months.

Sciurus variabilis (Variable Squirrel) 2 73 75

S. variegatus (Red-bellied ,, ) 13 36 129

S. vulgaris (Common iu iog tah 6 20

S. vulpina (Vulpine oe ULO 36 85

AGA MS. SNe ETUIS Sodeascossontageenace 24 9°8 4]

X..erythropus (Red-footed Ground 7 8 19

Squirrel).

AX. getulus (Getulian Ground 11 10°5 4]

Squirrel).

AX. setosus (Bristly Ground 6 10 15

Squirrel).

Tamias, whole genus ..........-.20+++- 16 26 84

T. asiaticus (Asiatic Ground Squirrel) 3 8 23

7. lateralis (Side-striped_,, ae 1 All 21

7. striatus (Chipping Squirrel) ...... 12 3] 84

Pteromys, whole genus ......-.....+.+--- 5 61 163

P. albo-rufus (Red-bellied Flying- 1 163 163

Squirrel).

P. inornatus (Large Red Flying- 1 2 2

Squirrel).

P. leucogenys (White Cheeked 1 101 101

Flying-Squirrel).

P. magnificus (Red-bellied Flying- 1 36 36

Squirrel).

P. oral (Large Brown Flying- 1 3 3

Squirrel).

Sciuropterus, whole genus ............ 14 15 61

S. fimbriatus (Grey Flying-Squirrel). 11 13°7 61

S. layardi (Layard’s ,, Ona eng 1 1 1

iS. volucella (American ,, sO ne) Gor ne 30 Be

= volans.

Spermophilus, whole genus ............ 24 17 132

S. citillus (European Souslik)... 5 74 iit

S. grammurus (Line-tailed ,, )... 1 119 119

S. mexicanus (Mexican 2 : 3 A7 S52

S. mongolicus (Mongolian ,, _).. 4 10:7 22

S. tridecimlineatus (Thirteen-str iped 11 5D 22

Souslik).

Arctomys, whole genus.................- 23 64 164

A. caudatus (Long-tailed Marmot). 1 15 15

A. himalayanus (Himalayan ,,_) 1 79 79

A. hodgsont (Hodesonis mos 6 ae 73°5 74

A.marmotia (Alpine ia ee all) 82 164

A. monax (Quebee Pai) 9 aye 116

Cynomys ludovicianus (Prairie ,, ) 48 30 98

JT am unacquainted with any published observations as to the

= 6X ON

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 447

longevity of Squirrels and other members of the Sciuride, and it

was a surprise to find that a Squirrel had lived over fifteen years,

a Flying-Squirrel over thirteen, a Souslik eleven, and a Marmot

over thirteen. It first occurred to me that as these animals

have not an individuality that is very obvious, mistakes with

regard to individuals might have confused the records. Such an

explanation, however, does not cover the facts; it would apply at

least equally well to the Common Squirrel, the 77 examples of

which showed no case of great longevity and a very low average,

and it could not possibly account for the many cases of high

longevity, where only two or three examples are recorded through-

out the period of thirty-three years. Wemust takeas established

the striking fact that squirrel-like animals have a very high

potential longevity in proportion to their size and in comparison

with other vegetable-feeding forms, as the latter are on the whole

relatively short-lived. The average duration and therefore the

viability of the group appears to vary much ; some species are

almost uniformly unsuccessful, whilst others show a distribution

of durations very close round the average. Taking the group as

a whole, however, it has shown a relatively high viability just as

it has a relatively high maximum longevity. It is particularly

interesting to reflect that during the greater part of the period to

which the figures relate, most of these squirrels were housed in

outdoor cages freely open to the air, with no artificial heating,

but with small nesting-boxes as retreats.

No. of Av. dur. Max. dur.

CasTORIDS. Indiv. in months. in months.

Castor fiber (Huropean Beaver) ... 4 A-2, 7

C. canadensis (Canadian i wee fal) 403 130

The great difference in viability between European and Canadian

Beavers is remarkable. The average of the latter is relatively

low on account of the large proportion that lived only a few

months ; if two or three months were survived these beavers usually

lived some years. The number of European beavers within the

period is small, and it may well be that it included no healthy,

nearly adult individual.

No. of Av. dur. Max. dur.

Myoxip2. Indiv. in months. in months.

Mayonus, Whole GENUS 222... seen 23 14:6 35

Myoxus dryas (Oak Dormouse) ...... 6 12°5 30

M., glis (Fat +3 Dhvmaercaer 3 14:3 34

M. quercinus (Garden _,, Ve ee ae 14 15:5 35D

Muscardinus avellanarius (Common 17 oD 13

Dormouse).

The relative low viability of, or failure with, the common

English Dormouse is remarkable. A high longevity is not to be

anticipated for dormice, although I am unaware of any recorded

448 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

observations. Throughout the period in question, these animals

were kept in a badly ventilated house, usually kept hot.

No. of

MuRID&. Indiv.

Hydromyschrysogaster(Golden-bellied 6

Beaver-rat).

Gerbillus, whole genus ...............--- WA

G. cgyptius (Lesser Egyptian 39

Gerbille).

G. campestris (Field Gerbille) ...... 2

G. indicus (@inidian es ceseccc: 15

G. longifrons (Long-fronted Gerbille) 79

G. pygargus (White-rumped ,, ) 2

G. pyramidum (Large Egyptian,, ) 14

G. robustus (Robust seville saneny

G. shawt (Shaw’s i ) 28

Psammomys obesus (Fat Sand-Rat) . 1

Georychus capensis (Cape Mole-Rat). 1

Cricetus frumentarius (Common 12

Hamster).

Cricetomys gambianus (Gambian 8

Pouched Rat).

Mois, WHOIS BOWS G4506504s00doa0s40an0000 64

M. arboricola (Sydney Bush-Rat) ... 3

M. bandicota (Bandicoot Rat) ......... 2

M. barbarus (Barbary Mouse) ...... 20

M. blanfordi (Blanford’s Rat) ......... 3

M. cervicolor (FKawn-coloured Mouse) 2

M.exulans (Pacific Mouse) ......... 9

M. minutus (Harvest Mouse)......... 10

DU RiKonagony (U8) vole 1 tun) sadeoonpesecacs 13

M. sylvaticus (Long-tailed Field- 2

Mouse).

Tsomys variegatus (Varied Field-Rat)

Acomys hunteri (Hunter's Spiny 11

Mouse).

Arvicola, whole genus ............0..00. 14

A. agrestis (Short-tailed Vole)...... 7

A. amphibius (Water Vole) ............ 5

A poratenses a(banl= Wiole)eren teccer: 2

Myodes lemmus(Norwegian Lemming) 5

Fiber zibethicus (Musquash) ............ 7

Ay. dur.

in months.

Max. dur.

in months.

The only published statement regarding the age of mice with

which I am acquainted, is Metchnikoff’s assignment of “ five or

six” years as their limit. It may be supposed that the larger

forms have a higher potential longevity. The maximum duration

in the record is nearly 7 years in the case of an Egyptian Field

Rat. Comparatively few of the maxima approach this figure, and

the relation between the maximum and average durations is fairly

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 449

close. As in proportion to their size these rodents have rather a

high viability, it must be supposed that the conditions of detention

in the interior of a house are not very unfavourable to them.

SPALACIDH. ae ia eee aie

Rhizomys badius (Bay Bamboo-Rat). 5 17-4 4]

Dipropip&.

Dipus egyptius (Egyptian Jerboa)... 48 188 56

D. hirtipes (Hairy-footed Jerboa)... 30 16 36

Dipodillus simoni (Simon’s Dwarf 3 20 23

Jerboa).

D. campestris (Dwarf Jerboa) ......... if 12 12

Alactaga indica (Indian Jerboa) ...... tL 14 14

A, decumana (Persian Jerboa)......... 2 10°5 13

If Jerboas can be compared with rats and mice, the record of

nearly five years for an Egyptian Jerboa is what might be ex-

pected, but the maxima for most of the species and the average

durations show a low viability. The Jerboas have always been

kept in a warmed house.

OcropontiDa. Ty erie sn aa

Octodon cumingi (Cuming’s Octodon). 35 24 62

Ctenodactylus gundi (Gundi Rat) ... 2 DD 7

Myopotamus coypu (Coypu) ......... 39 30 121

Capromys brachyurus (Short-tailed 3 1] 15

Capromys).

C. pilorides (Fournier’s Capromys) ... 5 36 116

Aulacodus swindermanus (Ground- 2 10°5 16

Rat).

Some of the Octodontide are relatively large rodents and the

record of 10 years for a Coypu is not surprising. The viability

as shewn by the average duration is not high; in the case of the

Coypus, which have always been provided with access to the open

air, the average duration is lowered by high mortality amongst

young born in the Gardens.

Fysrricip# No.of Av.dur. = Max. dur.

7 Indiv. in months. in months.

a sinia WHOLE eS eMUS ya)saa- <meta 34 89 245

H. cristata (Crested Porcupine) ... 26 70°9 245

HT. javanica (Javan Vie 22 123 181

Hf. longicauda (Sumatran ,, Diners) 2 137 163

Atherura africana (African Brush- 8 26:3 AZ

tailed Porcupine). z

A. fasciculata (Indian Brush-tailed ,, ). 2 29 Th5O

Erithizon dorsatus (Canadian Por- 3 18°3 32

cupine).

Sphingurus, whole genus ............ 18 23 107

450. DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Av. dur. Max. dur.

Indiy. inmonths. in months.

Sphingurus insidiosus (Guianan Tree 1 40 40

Porcupine).

S. mexicanus (Mexican Tree Por- 1 ia 7

cupine).

S. prehensilis (Brazilian Tree Por- 9 15 AT

cupine.

S. spinosus (Spiny Tree Porcupine). 1 1 1

The Porcupines are another set of animals regarding the lon-

gevity of which I am unaware of published information, and it is

most interesting to find that they attain considerable ages, the

record of over twenty years for a common porcupine being re-

markable for, a relatively small vegetable-feeding animal. The

high average duration (over 7 years) of the whole genus Hystrix

shows that these animals have a high viability. The contrast

between the records of maximum and average durations for the

genus Hystrix and those for other genera 1s extremely interesting.

In the period in question, the species of Hystriz have been given

free access to the open air and unheated shelters, whilst examples

of other genera have been for the most part kept inside artificially

heated houses.

Capt. Stanley Flower’s record cases are not so high.

No. of Av. dur. Max. dur.

CHINCHILLID. Indiv. inmonths. in months.

Chinchilla lanigera (Chinchilla) ...... 38 30 84

Lagotis cuviert (Cuvier’s Lagotis) ... 1 18 18

Lagostomus trichodactylus (Viscacha) 30 32 81

DaAsYPROCTID.

Celogenys paca (Spotted Cavy) ...... 35 22 92

Dasyprocta, whole genus ............... 101 31 109

Dcisonoly (UNESUO IN) ceoacesobe esac 4 33°D 61

D. agutr (Golden Agouti) ......... IK) Bid 101

IDs CHIR AN Nyy INRUBUS N opi )). Bhs oeS bce 3 94 109

D. cristata (West Indian Agouti) 29 27 99

D.isthmica (Central American ,, ) 15 dl NG

D. mexicana (Mexican Bee Ve 11:5 14

D. prymnolopha (Hairy-rumped ,, ) 20 32 108

D. punctata (Punctated EN) Tuma) 28 84

CavuDs.

Dolichotis patachonica (Patagonian 18 30 93

Cavy).

Cavia flavidens (Yellow-toothed 1 10 10

Cavy).

(Ob sada (Sows CRNA) coccesaussvaces-u: 4 28 48

Cerodon rupestris (Rock-Cavy) ...... 4 14 40

Hydrocherus capybara (Capybara)... 18 33 114

Pras Na RarRO RW gg

oye a a

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 451

The Agoutis and Cavies are another instance of small Rodents

which come from various kinds of climates, but which have been

treated as outdoor animals and show a relatively high maximum

and average longevity.

Vi 7 7 ae .

Lepontp.®. ary btn Si main

De Hus, WHOLEIBENUS soscoccedsasccesees cs 23 13 fal

L.. americanus(American Hare) ...... 3 2 2

L. brasiliensis (Brazilian ad eee 3 6°3 10

L. ewropeus (Common Sn ree 10 20 71

L. nigricollis (Black-necked ,, ) ...... 1 1 1

L. sylvaticus (Wood Gat) eee 4 13 41

L. variabilis (Varying a ete 8 2 6:5 12

Metchnikoff has set down ten years as a great age for a rabbit,

and hence the record duration of nearly seven years for the

common hare isnot high. The very low average duration shows a

poor viability. As there would seem to be no special difticulty in

providing Leporide with conditions suitable to them, there is

probably some special reason for their low viability in captivity.

It is a remarkable circumstance that many of the most common

wild animals of Europe are difficult to keepin captivity. I think

it is quite possible that the fear of man which such animals have

acquired in their wild state, almost as a necessary condition of

their existence, by continuing to act after captivity, shortens the

life, often by accident, often by reaction of the mental state on

the general health. Animals from remoter parts of the world

have not an acquired intolerance of man to lose.

Summary of Rodentia.

It is unlikely that animals in captivity reach the potential

longevities of their races, and it is anew and somewhat surprising

fact that Rodents live so long as these records show. ‘There is

only a rough correspondence between maximum durations and

size, the figures amongst Rodents ranging from 20 years in the

Porcupine, 15 in some Squirrels, 13 in Marmots, 11 in Sousliks,

9 in Agoutis and Capybaras, down to 3 in Dormice. Metchnikoff

has shown (fom. cit.) that on the whole there is an inverse pro-

portion between the relative capacity of the large intestines and

the duration of life, animals with a capacious hind-gut usually

being shorter-lived in proportion to their size than those in which

the hind-gut is reduced. Rodents certainly present a marked

exception to his general rule, for in their case the hind-gut and

cecum are relatively very long and very capacious. The relatively

high viability of Rodents is another remarkable fact, and is most

striking in comparison with Insectivora. Their high specific

longevities and viabilities may be associated, I think, with the

452, DR. P. CHALMERS MITCHELL ON LONGEVITY AND

fact that they are a successful group, with remarkable power of

adaptation to different environments.

The most important practical result is the very striking series

of contrasts between the average and maximum durations of those

animals which have had free access to fresh air and of those which

have been deprived of such access by detention in heated houses,—

a contrast that seems independent of the natural habitat of the

animals in question.

‘ No. of Ay. dur. “Max. dur.

HYRACES. Indiv. in months. in months.

Hyrax capensis (Cape Hyrax) ...... 10 15-2 50

H., dorsalis (Dorsal acetone 2 18°5 36

T can find no other records which would throw light on the

potential longevity of these animals, the zoological position of

which is very doubtful. It is impossible to tell their ages

if they are adult on arrival, and therefore the maximum

duration of just over four years gives no reliable indication of the

potential longevity. As these animals have a most complex and

capacious hind-gut, and as they are vegetable-feeders, a high

viability or great longevity is not to be expected, according to

Metchnikoff’s views. In the period covered by these figures, the

Hyraces were almost invariably given access to open air.

Capt. Stanley Flower’s figures are lower.

‘ No. of Av. dur. Max. dur.

PROBOSCIDEA. Tin in MOnEKe. in anootite.

Elephas africanus(African Elephant) 1 101 101

E. indicus (Indian 4 as 196 290

Metchnikoff has already pointed out that there is no exact

basis for the popular ascription of very high potential longevity

to these animals. It used to be thought that they could live for

several centuries. Flourens, using his formula based on the age

at which the epiphyses unite with the long bones, set down 150

years as the limit of their age. Indian natives ascribe a limit of

from 80 to 150 years. On the other hand, the official list of the

Indian Government, cited by Brehm, shows that of 138 elephants

only one lived for more than 20 years after it had been purchased.

Twenty to twenty-five years’ duration in a menagerie is regarded

as good. The record in this list of about 24 years has been far

surpassed by a female Indian Elephant, brought as a young

animal by the Prince of Wales from India in 1876,and still alive

and apparently in quite good condition in our Gardens, although

its age is certainly over 34 years. I agree with the general con-

clusion of Metchnikoff, that elephants, in proportion to their

size, have neither a high potential longevity nor a good viability,

not much more than a hundred years being the probable age limit,

and twenty to thirty years a fair average duration.

aS ee

en eo) a a

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 453

UNGULATA.

1. Perissodactyla.

RHINOCEROTID#. the cane ae

Rhinoceros, whole genus. ...............+++ 10 110 342

f. bicornis (Two-horned African Rhino- — 1 271 271

ceros).

R. lasiotis (Hairy-eared Rhinoceros) ...... 3 132 342

R. sondaicus (Javan 5 Visor 1 130 130

RF. sumatrensis (Sumatran __,, )) ade 4 5 AL?

Ro unicorns (Indian Rte crth) bartesk 1 281 281

Rhinoceroses, like Elephants, are large vegetuble-feeding

mammals, with in proportion to their size a relatively low

potential longevity and viability. The maximum duration in

this record, nearly 29 years for a so-called ‘“ Hairy-eared”

example, has since been surpassed by an Indian Rhinoceros which

died after a duration of 40 years in our Gardens, and probably

therefore at an age of at least 42 years. The number of

examples in the record is not sufficient to make averages useful,

but these and information I have received elsewhere make it

clear that the viability is not good, and probably lower in the

case of the African species.

No. of Avy. dur. Max. dur.

TAPIRIDA. Indiv. im months. in months.

Tapirus americanus (American Tapir). 138 43 188

LS iG (Nibeive Mle yerie)) spa cesoookeenar 2 4 5

The record of over 15 years for an American Tapir no doubt is

under the potential longevity of these animals, but I can find no

reliable information from other sources. The very low figures for

the Malay Tapir refer only to two individuals, and there is no

sufficient reason to suppose that the Asiatic species is less viable

than the American. It must, however, be inferred that Tapirs

in regard to their size are short-lived creatures with low viability.

No. of Av. dur. Max. dur.

Kquips. Indiv. in months. in months.

Biggs wit@le Sens.) eas ascet ea eee 21 116 346

Jd), (OS TOWOOUIS (UEGERNG)) dao epenacd sore Skene 1 220 220

LE. hemippus (Asiatic Wild Ass)......... 3 82 169

(=hemionus indicus).

Li wOmegera (Onsen) cosa. ce ok nenee dance 2 198 219

LE. somalicus (Somali Wild Ass) ...... 1 16 16

Li. teniopus (African Wild Ass) ...... B 175 232

EL .guaggon (@Qiwaeeal he. te... .ensem 1 346 346

E. burchella (Burchell’s Zebra) ......... 7 98 268

EH. grevyt (Grevy’s Zebra) .........2.6505 2 6 10

L. zebra (Mountain Zebra) ............ 1 46°5 62

It is well-known that the usual duration of life in horses is from

15 to 30 years, and somewhere about 50 is probably the potential

Proc. Zoot. Soc.—1911, No. XXX. 30

ADA. DR. P. CHALMERS MITCHELL ON LONGEVITY AND

longevity of the group. The record of a 29 years’ duration in

the case of a Quagga is therefore quite good, and if it be

remembered that the figures include a number of newly imported

and young animals, the average duration and viability are both

good. On the other hand, they are in accordance with Metchni-

koff’s views as to the relatively low viability and longevity of

animals with capacious hind-guts, for they are not high in

proportion to the size of the animals and their ready submission

to captivity.

Summary of Perissodactyla.

These animals, in consideration of the ease of procuring suitable

food, do not afford evidence of high potential longevity or of great

viability im proportion to their size.

2. Artiodactyla.

No. of Ay. dur, Max. dur.

Bovin. Indiv. in months. in months.

Bosy umd rents; (Ae WW) if sesh cae ts eca 2 20 64:5 230

Bos taurus (English Domestic White 13 32 145

Ox).

Bison americanus (American Bison)... 9 70 129

B. bonasus (European |), 0)=-2 at 14 14

Poephagus grunniens (Yak) .........-+. 4) D6 107

Bibos fjrontalis (Gayal) ............-...-. D 138 188

JB. qremnpons (Clare) ceacecobe cosoessesccoees 1 30 30

Bubalus cequinoctialis (EB. African 2 aye 90

Buftalo).

B. buffelus (Indian Buffalo) ...........- 1 20 20

B. caffer (Cape PL) Sa ea 2 118-5 175

Anow depressicornis (AN0&) .........45 2 63°5 126

All Bovanests. Caer. s e. . ceeaiee Seat. 61 63°3 230

According to Metchnikoff, the duration of life in cattle is very

short considering their size. The potential longevity is probably

not much over 30 years; at 5 years old they begin to show signs

of age and at from 16 to 18 they lose reproductive powers. The

records quoted above are wholly in accordance with this estimate ;

the maximum duration of nearly 20 years 1s good, and the average

duration is what might be expected reasonably. Throughout the

period in question, the cattle were kept in unwarmed shelters,

with access to the open air.

Capt. Flower has no high records.

No. of Av. dur. Max. dur.

BUBALIN. 3 Indiy. in months. in months.

iB. cacnion (ElarieWees) eeeten--eeneeee eee: 3 65 85

J, troinay (Mores ate) OOS), sscdnscccscoses sue 3 5D 86

Damaliscus albifrons (Bless-bok) ...... D 120 133

DS pygarges | sone Dos) a -eeee eee eee 2 28 55

Connochetes giw (White-tailed Gnu). 4 102°5 164

Adlsipulloalimes: isaac een eae 19 83 162

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 455

The records of Bubaline Antelopes correspond closely with

those of cattle. From their smaller size so high a potential

longevity is not to be expected, whereas, on the other hand, the

higher average duration is probably due to the absence of young

calv es, the mortality amongst which lowered the record of cattle.

CEPHALOPHINA. Twi. reais ae

Cephalophus, whole genus ............... 49 23 109

CO. coronatus (Crowned Duiker)......... A 1] 20

C. dorsalis (Bay Pea Tae Fi 7 26 (D

C. grimmi (Common 5 Aa) eka 13 eT, 107

CO. maxwelli (Maxwell’s ,, _)......... 10 34 109

C. monticola (Blue ys Gea 9 14-9 A3

C. nigrifrons (Black- outed Duiker) 1 3) 3

C. rutilatus (Red-flanked ee ne 27 38

C. sylvicultria (Yellow-backed nl 3 3

Tetraceros quadricornis (Four-horned 19 28 66

Antelope).

The maximum and average durations of the Cephalophine

Antelopes are rather lower in proportion to their size than those

of the Bubaline Antelopes.

There has been omitted from the

tables, however, a considerable number of examples that lived less

than a month in the Gardens.

Max. dur.

NEOTRAGINA. Noten yy, dur.

Indiv. in months. in months.

Oreotragus saltator (Klipspringer) ...... 1 46 AG

Raphicerus melanotis (Grys-bok)......... 2 6:5 10

R. tragulus (Stein-bok) «1........0:.0.062. 2 3 4

Ourebia nigricaudata (Gambian 3 40, 105

Ourebi).

The number of individuals is too small to make inferences

useful.

CERVICAPRINE. No. of _ Av. dur. Max. dur.

Indiv. inmonths. in months.

Cobus ellipsiprymnus (Common Water: 3 39°3 47

buck).

C. unctuosus (Sing-sing Water-buck). 4 100 183

Cervicapra isabellina (Isabelline 2 19 36

Antelope).

C. bohor (Bohor Antelope) ............... 2 43 83

ANTILOPINA.

Gazella, whole @enus /...75...-0......-.. 118 24-4 112

G. arabica (Arabian Gazelle)....c2...... 20 18 71

G. bennetti (Indian Rega) oes rat SE 11 10:8 57

Grcuvier’ (CuvieE Ss sy) ) ese ceeae.. ] 3 3

G.-dorcas (Hgyptian ,, ).:....20078 27 22°7 94

30*

456 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Av. dur. Max. dur.

ANTILOPINE (cont.). Indiv. in months. in months.

Gazella loderi (Loder’s Gazelle) ...... il 55 55

G. marica (Marica Per) eae 2 4] fal

G. mohr (Mohr’s a 1 58 58

G. muscatensis (Muscat ;, ) .-.... 8 18 45

G. rufifrons (Korin sia pele: cere 2, Ot 38

G. semmerringt (Scemmerring s 8 31 69

Gazelle).

. spekit (Speke’s Gazelle) ............ 2 :

. subgutturosa (Persian Gazelle)...... 26 33°D 112

Antilope cervicapra (Indian Antelope). t

The Gazelles in proportion to their size haye a good maximum

and average duration.

HIProTRAGINA.

ITippotragus equinus (Equine 1 123 123

Antelope).

H. niger (Sable Antelope) ............ Z 68 89

ORYGINE.

Ongmawiiole seems iy sanso ss: ee ceae eee Hp 15 83 216

O. beatrix (Beatrix Antelope) ......... 6 24 90

O. beisa (Beisa penn ts) een he ie. 4 129 WeT

Onlewconya (Welcony xp.) )\ere ee D 127 216

Addax — naso-maculatus (Addax 2 178 219

Antelope).

With exception of the Beatrix Antelope the members of this

fair-sized and handsome group display a remarkable longevity,

18 years being the record, and a high average viability.

No. of Ay. dur. Max. dur.

TRAGELAPHINS. Indiv. in months. in months.

Orcasieanna (Bland )\\ Reece heer eee 16 71 118

Strepsiceros kudu (Kudu) ............... 6 35 94

Se mevenbisn lesser mkGuciun iment Eere 2 44-5 80

Tragelaphus gratus (Pleasant Antelope) 2 - 42-5 78

T. scriptus (Harnessed Antelope) ...... 12 36 72

T. sylvaticus (Bosch-bok) ............... 4 49 82

Boselaphus tragocamelus (Nylghaie)... 9 71:4 146

The large Tragelaphine Antelopes have a proportionally high

maximum duration and their average duration shows that even

including the Kudu, which has been supposed to be specially

delicate in London, they have a good viability.

RUPICAPRIN A.

Rupicapra tragus (Alpine Chamois)... 1 24 24

Nemorhedus goral(Goral Antelope)... 1 212 212

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 457

~The most striking feature about these records of Antelopes is

that in proportion to their size, they appear to have better lives

than the cattle, Within the group there is a fairly close

correspondence between size and duration, but a few species

such as the Beatrix and the Chamois are specially delicate. The

conditions under which these animals have been kept vary as to

the provision of artificial heat, most of the Gazelles having been

least well provided with it, but in nearly every case there has been

access to the open air.

Capt. Flower’s records of maximum duration are specially

interesting, as it is to be expected that the climate of Hgypt

would be specially favourable to Antelopes. His highest figures,

however, do not differ much from ours; 8 years for a Cobus

defasst, still alive, over 11 years for a Dorcas Gazelle, 10 years

for a Seemmerring’s Gazelle and for a Sabre-horned Oryx.

No. of Ay. dur. Max. dur.

CAPRIN 2. Indiv. im months. in montlis.

Capra; whole venus MNT. JAE Ae Ge 50 50 166

C. cegagrus (Grecian Ibex)............... 5 59 157

C. caucasica (Caucasian Ibex) ......... 2 54° 78

C. hircus (Angora Goat) ............65. 18 53°5 166

C. hispanica (Spanish Ibex) ............ i 32 32

CO. ibex (Alpine yal) See. 5: 2 103 103

C. megaceros (Markhoor) ............... 15 5D 133

C. nubiana (Nubian Ibex)............... 6 24 101

CG; sinaituca, (Simatic, F)\)a:y. eee: 1 45 45

Hemitragus jemlaicus (Thar) ............ 10 51 143

Orisrawihole emus caster each: acs viaen 68 46 231

O. blanfordi (Blanford’s Sheep)......... 1 46 Ab

O. burrhel (Burrhel Wild Sheep) ...... 21 46°7 193

O. cycloceros (Punjab ,, 8 i ae sre 6 51 3a

O. hodgsoni (Great Tibetan Sheep) ... 2 20 31

O. musimon (Mouflon) .................. 15 50 231

O. tragelaphus (Barbary Wild Sheep). 21 43 134

Orgingnied, (Wire) hi eters! eats 340 ee 15 28

Sheep and goats are supposed to have a shorter life than

cattle. Metchnikoff, quoting Grindon, assigns 14 years as an

extreme age for domestic sheep, 12 an old age, and 8 to 10 as

the time when senile changes are expected. These records show

that wild species of sheep and goats may attain great ages, a

goat having lived about 14 years and a Mouflon over 19 years in

the Gardens. The average duration is lowered by the number of

lambs the life of which was not good, and if this allowance be

made, it is clear that the sheep and goats have a fairly good

viability. They are of course kept with free access to the open air.

Capt. Flower gives no higher records,

458 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Ay. dur. Max. dur.

ANTILOCAPRIDE. Indiv. in months. in months.

Antilocapra americana (Pronghorn) win gO 9 30

Pronghorn Antelopes are notoriously delicate in captivity, not

only in London, but in other zoological collections. The records

afford no information as to the specific longevity, and only

confirm knowledge as to the low viability of these animals.

GIRAFFID&.

Giraga, whole genus..........--...-..-.- 10 101-9 229

It is to be expected that Giraffes, in correspondence with their

large size, should have a relatively great specific longevity com-

pared with other ruminants. The record, a duration of over

19 years, is just surpassed by that of a Zebu, and whilst it is

fairly high, is certainly not high in proportion to the great size

of the animal. The average duration shows a fairly good viability.

Giraffes have always been given artificial heat in cold weather, and

for the most part have been allowed access to the open air only

in summer.

Capt. Flower records a Giraffe still alive after 8 years’ duration.

CERVIDA.

No. of Ay. dur. Max. dur.

MOoscHIN#. Indiv. in months. in months.

Moschus moschiferus (Musk-deer) ...... 4 26 65

Musk-deer have shown a low viability. There is some difficulty

as to their food in captivity, as their natural diet of twigs and

buds is difficult-to supply; they are animals capable of surviving

extreme exposure to cold, and in captivity have probably been sub-

jected to too much artificial warmth.

No. of Av. dur. Max. dur.

CERVINA, Indiv. in months. in months.

Cervulus, whole genus .............6..+: 14 71 126

C’. crinifrons (Hairy-fronted Muntjac). 1 73 73

CO. lacrymans (Crying Muntjac) ...... 1 126 126

C. muntjac (Indian SU) ae He 7 39 96

C’. reevest (RESTESIS 55) sanosc 5 105 12a

Hlaphodus nuchianus (Michie’s Tufted 9 16 46

Deer).

Cercus, whole genus 58. See. 173 68°8 243

Red-deer group.

Cervus elaphus (Red Deer)............... 17 69 191

C. kashmirianus (Kashmir Deer) ...... 1 148 ets

C. maral (Bersiaii ca) ceed 3 98 172

C. luehdorfi Gailidortis sas) eae 2 525 79

C. canadensis (Wapiti etic 0 156°9 243

Oi a a i

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 459

No. of Av. dur. Max. dur.

Sika SOE) E Indiv. inmonths. in months.

Cervus sika (Japanese Deer) ............ 39 52 160

C. mantchuricus (Mantchurian Deer). 4 116 170

C. tagvanus (Formosan Oa en ee 95 149

Axis group.

Osarisi(Agiis Weer) in ayste sca sebees s-. 27 56 158

Sambar group.

C. aristotelis(=wnicolor)(Sambar Deer). 17 48 142

CO. hippelaphus (Rusa _ Deer)......... 3 26 ai

C. moluccensis (Molucca _,, )......... 12 69 219

C. philippinus (Philippine ,, )......... 3 5D 56

C. kuhli (Kuhl’s AY 0) aenaneeee 2 37 66

C. alfredi (Prince Alfred’s ,, )......... 5 51 93

C. porcinus (Hog Ai) Senate 16 60°5 119

Swamp Deer group.

O. duvaucelli (Barasingha Deer)... 1 220 220

C. schomburgki (Schomburgk’s ,, ).... 1 157 157

C. eldi (Panolia se eee Caius) 58 91

C. davidianus (Pére David’s Deer) ... 3 49 92

Dama mesopotamica (Persian Fallow 8 60 124

Deer).

D. vulgaris (Common Fallow Deer)... 8 42 120

Alces machlis (Moose)..........5........+5 6 32:7 2

Hydropotes inermis (Chinese Water- 3 70 129

Deer).

Capreolus caprea (Common Roe-deer). 9 24 52

C. pygargus (White-rumped Roe- 1 6 6

deer). )

Cariacus, whole genus ...............-.- 83 31 146

C. campestris (Pampas Deer) ......... 6 12 24

C. gymnotis (Naked-eared Deer)...... D 14:5 40

C. leucurus (White-tailed ,, )...... 1 69 69

C. macrotis (Mule sate) eet 24° 49 146

C. mexicanus (Mexican Ae or 18 31 91

C. nemorivagus (Wood Brocket)...... oo 17 32

C. rufus (Red bw tire 6 13} 35

C. virgimanus (Virginian Deer)...... 16 27 81

Furcifer chilensis (Gemul ,, )...... 1 131 13

Pudua humilis (Pudu » iehaes + 31 108

Rangifer tarandus (Reindeer) ......... 1] 4] 116

Stags are supposed to be long-lived animals, and Flourens has

set down 30 to 40 years as the duration of their lives. The

longest record. on this list, over 20 years in the case of a Wapiti,

does not lead one to infer that in proportion to their size-the.

460 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

potential longevity of Cervide differs much from that of other

ruminants. Within the group, there is the usual rough corre-

spondence between duration and size, the larger animals reaching

greater ages and being hardier than the smaller animals; the

records of average duration are lower, in some cases (¢. g. Sika,

Axis and Red Deer), reduced by breeding, the mortality amongst

the young being heavy. Some kinds of Deer have a specially

low viability in captivity, as for instance the Moose, most

probably on account of the difficulty of supplying it with

natural food, whilst all Deer of the genus Cariacus except the

Mule Deer appear to be naturally delicate. The vast majority

of the deer have been kept under similar conditions, with

unwarmed shelters and free access to the open air.

Capt. Flower’s highest record is that of an Axis-deer, still alive

after over 9 years’ duration.

No. of Av. dur. Max. dur.

TRAGULIDE. Indiv. in months. in months.

Uraguics wo leye ens) oe ees creer cre 28 11-4 50

T. javanicus (Javan Chevrotain) ......... 10 9:9 43

T. meninna (Indian Fs eee y 12 33

T. stanleyanus (Stanley’s ,, Dy octane 9 12 50

Hyomoschus aquaticus (Water Chevro- 11 4 lel

tain).

Nothing is known as to the potential longevity of Chevrotains,

and the record of over four years would appear to be considerably

lower than what might be anticipated for these animals. The

viability in captivity is very low, and the records of average

duration would be much lower if a considerable number of

examples that lived for less than a month had been included.

These animals have always been assumed to require special

protection from cold and have been kept in warmed houses with-

out access to the open air.

No. of Av. dur. Max. dur.

CAMELID. Indiv. im months. in months.

UGG, WANON® GEWS anchoasoeeoncoss00adeeon 14 69 187

aiemanacos| (ciumanaco)i ieee 6 72 187

Ibs FOGHGOS (PN DENCE) cau boceoayaecceeecessog esd 2 9 14

th, jagpuiming (WMENIIE)) 4.64. sochonsceose9000r 6 95 156

Camelus dromedarius (Dromedary)...... 6 44 145

O. bactrianus (Bactrian Camel) ......... 5 110 204

Flourens has set down the life of a Camel as reaching to

40 years, but the longest record amongst the Camelide is only

17 years. A Llama that died at the Gardens recently, after a

duration of 15 years, showed plain evidence of senile degeneration.

The figures of average and maximum duration show that the

Camelidee, notwithstanding their size, and the absence of difficulty

in providing appropriate food and accommodation, are not long-

lived animals.

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 461

No. of Av.dur. Max. dur.

HippororaMip&. Indiv. in months. in months.

LTippopotamus amphibius (Hippo- 3 290 335

potamus).

The maximum duration of nearly 28 years has been surpassed by

a female Hippopotamus, born in the Gardens and which recently

died at the age of 39 years. The viability of these animals is

good, if they are sufficiently protected from cold in winter, but it

is to be noticed that if the maximum ages attained give any

indication of the potential longevity, the Hippopotamus, in pro-

portion to its size, cannot be regarded as a long-lived animal.

No. of Av. dur. Max. dur.

PHACOCHGRIDE. Indiv. in months. in months.

Phacocherusafricanus(Atlian’s Warthog) 1 24 24

P. ethiopicus (AMthiopian Warthog)...... 6 5D 131

Su.

SeUsmWNOle BeMUS: eek. kee nem eee 25 DD 234

Sus andamanensis (Andaman Swine) ... 2 134 147

S. cristatus (Indian Wild Swine) ......... 2 54:5 89

S. leucomystax (White-whiskered Swine) 5 65:8 158

S. papuensis (Papuan Pig) .... .......... 1 74 74

WAISCHOPO) (NW UGS WAC), ean aaen santyteg. «ne. 15 40 234

Porcula salvania (Pigmy Hog)......... .. 5 48 90

Babirussa alfurus (Babirussa) ............ 6 A] 126

Potamocherus africanus(Southern River- 5 86 176

Hog)

P. penicillatus (Red River-Hog) ......... 6 4 ll

Dicotyles labiatus (White-lipped Peceary) 5 Ol 112

D. iagacu (Collared Peccary) ............ 20 22°9 162

The maximum duration of nearly 20 years for one of the Wild

Swine corresponds with the limits that various authors have

assigned to such animals. The viability is not good, and the

average would be much lower if a number of animals whose

duration was less than a month had not been omitted.

Capt. Flower’s highest records are that of a Wild Swine still

alive after 12 years’ duration, and a Babirussa alive after 9 years’

duration.

Summary of Artiodactyla.

The Artiodactyla contain animals of so many different kinds,

that we cannot expect to find a close correspondence between

size and longevity throughout the whole group, although on

the whole such a correspondence exists throughout the sub-

groups. The group as a whole is not notable for longevity in

proportion to size, and as the majority are vegetable-feeders with

capacious hind-gut and great tendency to intestinal putrefaction,

the general condition is in agreement with Metchnikofi’s theory.

Antelopes, sheep and goats, and deer appear in proportion to

their size to have better viability than cattle, giraffes and camels,

whilst on the whole the non-ruminants are hardier, in proportion

to their size, than the ruminants.

462 DR. P, CHALMERS MITCHELL ON LONGEVITY AND

CETACEA anp SIRENIA.

Very few examples are contained in the records, and the

duration of these was only sufficient to show the complete

unsuitability of the conditions and certainly afforded no indication

as to the general viability and longevity of these animals. There

is of course abundant evidence from other sources as to the high

potential longevity of Cetaceans, but I know of no definite

evidence with regard to Sirenians, which as vegetable-feeders

with complicated intestines might be supposed to have short lives

in proportion to their bulk.

EDENTATA.

= No. of Av. dur. Max. dur.

BRADYPODIDS. Tinie. im months in aoe

Bradypus tridactylus (Yhree-toed Sloth). 8 underl 1

Cholopus didactylus (Two-toed Sloth) ... 2 7 13

C. hoffmanni (Hoffmann’s Sloth)......... 14 29 133

I am ignorant as to any recorded information regarding the

potential longevity and viability of Sloths. Lydekker, in the

Royal Natural History, remarks on their power of recovery from

injury and capacity for enduring long periods of starvation,

correlating these qualities with general low organization, and

comparing the animals with reptiles. Certainly, however, it

cannot be assumed that creatures of low organization have a

relatively high potential longevity. As Sloths are rather small,

vegetable - feeding animals, with fairly bulky hind-guts, on

Metchnikoft’s theory, their duration of life and viability ought

to be low. The maximum record of over 11 years for a

Hoffmann’s Sloth is certainly high, but the average duration of

Sloths is extremely bad. The failure has been complete in the

case of the Three-toed Sloths, very great with the Two-toed,

whilst there has been a partial but very irregular success in the

case of Hoffmann’s Sloth. Sloths are nocturnal inhabitants of

tropical forests, and in captivity they have been assigned confined

quarters in warmed houses, without access to the open air.

Their quiescence by day probably has led to the need of air and

exercise at night being overlooked.

DASYPODIDA. a of Av.dnr. Max. dur.

ndiv. in months. in months.

Tatusia hybrida (Mulito Armadillo) ... 6 8:5 AQ)

Ts pebon (Reba Aucmneadalllo)) ae: -assenseeneeee: 16 7 4D

T. kappleri (Kappler’s Armadillo) ....... 2 53 D4

Dasypus minutus (Little Armadillo) ... 6 19 23

D. sexcinctus (Six-banded Armadillo) ... 6 $°3 19

D. villosus (Hairy Armadillo) ............ 31 33 157

NXenurus wunicinctus (Broad-banded 2 6 11

Armadillo),

Tolypeutes tricincius (Brazilian Three- 1 8 8

banded Armadillo).

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 463

Regarding the potential longevity of Armadillos, I am unaware

of any recorded information. They live on carrion, insects and

vegetables, and the hind-gut is of very moderate relative capacity.

The record of 13 years for a Hairy Armadillo seems good, but the

average duration is very much less than that. Throughout the

period in question most of the Armadillos were kept in the

interior of a badly ventilated and very warm room.

Capt. Flower records a duration of over 8 years for a Hairy

Armadillo.

No. of Av.dur. Max. dur.

MYRMECOPHAGIDE. Indiv. in months. in months.

Myrmecophaga jubata (Great Ant-eater). 10 46 172

Tamandusa tetradactyla (Tamandua Ant- 12 10°8 60

eater).

The records of 144 years for a Great Ant-eater and 5 years for

a Tamandua Ant-eater are very much in excess of the average

duration of these animals. They have always been carefully

protected from cold, and not allowed access to the open air. It is

plain that they are creatures of low viability compared with their

potential longevity, but that they have been subjected to very

unsuitable conditions.

No. of Av. dur. Max. dur.

MANID&. Indiv. in months. in months.

Mamistricuspis (White-bellied Pangolin). 1 1 1

WM. (Pholidotis) gigantea (Giant Pangolin). 1 1 1

The records only testify to the failure to keep these animals in

captivity.

ORYCTEROPIDS.

Orycteropus capensis (Cape Ant-bear) ... 4 52 LZ

O. ethiopicus (Aithiopian Ant-bear) ... 1 Le 17

The record of over 9 years for a Cape Ant-bear gives an

indication that these animals are able to live to a considerable

age. The difference between the maximum and average durations

shows the low viability of these animals under the conditions of

their captivity. In the period in question they have been kept in

heated houses without access to open air.

Summary of Hdentata.

The most notable feature in the records is their great variation.

The maximum durations are sufficient to deduce from them that

these animals have a fairly high potential longevity which varies

roughly with their size, and that their viability is low under the

conditions in which they have been kept.

464 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

MARSUPIALIA.

No. of

MAcROPODIDA. Tondlicr.

Macropus, whole genus|......)....2.....-.:- 200

M. agilis (Agile Wallaby) ...... 5

M. bennetti (Bennett’s ,, ) ...... 35

M. billardiert: (Red-bellied ,, yak derek ob

M. brachiurus (Short-tailed Wallaby) .. 15

M. derbianus (Derbian santa) een TKO)

M. dorsalis (Black-striped ,,_ )... 8

M. giganteus (Great Kangaroo) ......... 30

M. melanops (Black-faced Kangaroo)... 13

M. parryi (Parry’s Wallaby)............... ]

M. robustus (Great Wallaroo) ............ 15

M. ruficollis (Rufous-necked Wallaroo) . 6

M. rufus (Red Kangaroo):.::.............. 30

M. thetidis (Thigh-striped Wallaby)...... 4

M. walabatus (Black-tailed ,, _ )...... 12

Petrogale, whole genus A4

P. brachyotis (Short-eared Kangaroo)... 1

P. penicillata (Brush-tailed ,, ).. 15

P. xanthopus (Yellow-footed Rock 28

Kangaroo).

Onychogale frenata (Bridled Wallaby)... 12

O. unguifera _ (Nail-tailed SN) eo 6

Dorcopsis luctwosus (Mourning Kangaroo) 1

D. miillert (Miiller’s Kangaroo) ......... 1

Dendrolagus tivustus (Brown Tree Kan- 1

garoo),.

D. bennetti (Bennett’s Tree Kangaroo) . 4

“Epyprymnus rufescens (Rufous Rat- 19

Kangaroo).

Bettongia, whole genus .............0....00. 24

B. cuniculus (Tasmanian Rat-Kangaroo). 3

B. gaimardi (Gaimard’s _,, Ae eb)

SL. leswewrt (Lesueur’s _,, Hh aes 3

B. penicillata (Ogilby’s _,, 3

Potorous tridactylus (Common Rat-Kan- D

PHALANGERID#. gar00).

Dromicia nana (Dormouse Phalanger) . 4

SHON TONPOIS, WDONEY CXETONWIS Goodnocosossos +05 +06 48

P. australis (Yellow-bellied Phalanger). 2

P. breviceps (Short-headed ss Ve oy

P. sciureus (Squirrel-like 4 sped

Pseudochirus peregrinus (Ring-tailed 1

Phalanger).

Trichosurus, whole genus .................- 71

T. caninus (Short-eared Phalanger)...... 1

T. fuliginosus (Sooty his AO) ae 7

T. orientalis (Oriental ff Daas 1

T. vulpecula (Vulpine oh yee 6

Phascolarctos cinereus (Koala) ............

Ay. dur.

in months. in months.

30°2 166

21 43

30 12

12 38

16 4]

38 118

23 62

31 13

Bil 164

5 5

25 83

14 28

43:8 166

26 54

30 76

30 89

3 3

31 89

31 79

Bes} 10

1 2

7 7

15 15

19 30

18 43

35D 70

40 94

29 59

30 79

78 94.

Lay 87

13 16

26°5 64

AO 130

92°5 121

33 130

48 123

AL A

22 142

89 89

27 76

3 3

21 142

5°6 8

Max. dur.

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 465

It is impossible to separate, by any important characters, the

Kangaroos, Wallabies, Rock Kangaroos, Rat-Kangaroos, and

Phalangers. They form a series roughly corresponding with the

ruminants amongst higher mammals, and have similar habits

and food. The alimentary tract, although much simpler in

structure than that of ruminants, possesses a capacious hind-gut

and cecum, and they are theretore animals that might be

expected to have low potential longevity and poor viability.

The maximum duration in the group, nearly 14 years in the case

of a kangaroo, is more or less what might be expected, and on

the whole there is a rough correspondence between size and

longevity. The average duration is low compared with the

maximum duration, and it is a fair inference that the viability is

not good. Amongst the Phalangers, however, there are remark-

able exceptions, the maximum duration and the average duration,

in a number of cases, being unusually high in proportion to size.

In reflecting upon this I was disposed at first to distrust the

records ; in animals so little individualized it might well be that

mistakes had been made as to individuals, and hence that wrong

durations had been assigned in the records. Although this may

well have happened in some of the cases, it is not a sufficient

explanation. A period of 33 years is covered by the records, and

even where the number of individuals was largest, the number of

individuals alive at any one time was probably not more than 3 or

4 at most, and the chance of confusion was therefore not great ;

there are moreover cases of remarkable duration where the

number of individuals was so small as to make confusion prac-

tically or absolutely impossible,—-of two Yellow-bellied Phalangers,

for instance, one lived over 10 years, and one Short-eared

Phalanger lived over 7 years. It is clear that these animals have

a high viability. In the period covered by the record some of

them were kept in fairly roomy cages, but in an artificially heated

house without access to the open air. Others were kept in small

outdoor cages with no artificial heat. Unfortunately J cannot

now trace which individuals lived in these very different kinds of

quarters. Since the period in question several species have

thriven ina shed open to the outer air, without artificial heat,

but with protection from wind. Capt. Flower records a Great

Grey Kangaroo, a Black-faced Kangaroo, and a Wallaroo, each

alive after 10 years’ duration.

No. of Av.dur. Max. dur.

PHASCOLOMYID2. Indiv. in months. in months.

Phascolomys, whole genus...............-+- 13 74 212

P. latifrons (Hairy-nosed Wombat)...... 4 88 212

P. mitchelli (Common Sy ae 9 68 189

A record of nearly 18 years for a Wombat seems to show that

these animals have a high potential longevity in proportion to

their size, whilst their viability, as shewn by the average duration,

is good. Throughout the period, they were kept in an unheated

shelter with free access to the open air.

466 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

PERAMELID2. Indiv. in months. in months.

Peragale lagotis(Rabbit-eared Bandicoot). 8 19°8 32

Perameles obesula (Short-nosed ,, ). 3 SUnu 17

These little omnivorous and carnivorous marsupials show low

maximum and average durations. They havea capacious hind-gut

and ezecum in proportion to their size. They have been kept in

small cages in a heated house, without access to fresh air, and

have not done well.

No. of Av. dur. Max. dur.

DasyurID&. Indiv. in months.: im months.

Thylacinus cynocephalus (Tasmanian 7 43 103

Wolf),

Sarcophilus ursinus (Tasmanian Devil). 7 52°5 61

Dasyurus, whole genus ..........--...-..... 44 33 82

D. maculatus (Spotted-tail Dasyure) ... 5 37 15

D. viverrinus (Maugés Adc i) asta “eo 28°5 82

Compared with Carnivora of similar sizes, the maximum

duration of these carnivorous marsupials is not so high, sug-

gesting possibly a low potential longevity. On the other hand,

the average duration is rather better in proportion, which may

be evidence of a greater viability. The conditions under which

they have been kept are similar to those of true Carnivora: larger

forms like the Thylacine have been given unheated shelters with

access to the open air; the smaller forms, for the most part,

have been confined in artificially warmed houses.

No. of Av. dur. Max. dur.

DIDELPHYID ®. Indiv. in months. in months.

Didelphys, whole genus ......----...-..+.-+- 92 10°8 85

D. azare (Azara’s Opossum) .......--.6+ 4. 20 8 28

D. cancrivorus (Crab-eating Opossum). 31 13 85

D. cinereus (Cinereous ah Real 5 5

D. crassicaudata (Thick-tailed 9 SEEMED. 16 28

D. lanigera (Woolly it OF 66 19 53

D. murina (Murine Wits ia tay 62 1:5 9

D. nudicaudata (Rat-tailed Pt) 2 9) 14

D. opossum (Quica sor we) 9 1] 22)

D. philander (Philander Seiad A 4-7 9

D virginiana (Virginian am ys 12 8 16

The Opossums, which are omnivorous and carnivorous, display

low maximum and average durations, and so far as this evidence

goes would appear to have a relatively low potential longevity

and a poor viability. On the other hand, they have been treated

as animals requiring protection from cold at all costs, and have

been kept in artificially warmed houses without access to the

open air,

er a a

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 467

Summary of Marsupialia.

The records show in the first place that whilst within the

different divisions of Marsupials, there is frequently a close

correspondence between size and longevity, there is no such

relation if the group be considered as a whole. The maximum

duration was that of a Wombat which lived nearly 18 years in

the Gardens; next comes a Kangaroo with a duration of 14 years;

then some small Phalangers reaching nearly 12 years; then a

Thylacine reaching nearly 9 years. It is notable that much the

best records are in the case of animals which have had good access

to air; the chief exception, and this only a doubtful one, is that

of the Phalangers.

No. of Av. dur. Max. dur.

MONOTREMATA. Indiv. in months. in months.

Echidna hystrix (Echidna) .............:.. 4 1°5 5

The records are sufficient only to show the failure, in the period

in question, to provide suitable conditions for these animals.

They have always been kept in artificially heated houses.

SUMMARY OF MAMMALIA.

It is advisable to repeat here that the material on which this

communication is based, is defective in certain important respects.

No information was available as to the condition of the animals on

their arrival. I have tried to eliminate this source of error to a

certain extent, by omitting in most cases entries of animals that

had lived less than a month in the Gardens. Similarly, no infor-

mation as to the age of entrants wasavailable. It is obvious that

the heavy incidence of mortality in very young animals is an

important factor independent of the special conditions of captivity,

whilst the possibility of duration is naturally less in old animals

than in young adults. These defects probably have abnormally

lowered the records of average and of maximum duration, but to

what extent I am unable to guess. At the beginning of 1911,

I put into operation a system suggested to me by my friend

Mr. J. L. Bonhote, F.Z.S., and I hope that thirty years after this,

someone will be in a position to institute an enquiry similar to the

present communication to the Society, but based on more exact

data. In every case where it is possible, a card is prepared for each

new arrival at the Gardens; on the face is placed the name, and

mode and date of acquisition of the animal; on the reverse is

entered the first place of detention, the apparent condition of

health, and any indications of age (“very young,” “adult,” or

“old adult,” unless exact particulars are known); the card follows

the animal to each successive house in which it may come to be

placed ; the dates of such changes and any important events in

its history are noted; finally, at the Prosectorium, the date and

cause of death are entered, with any further data as to age that

may be available.

468 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

In the meantime, however, it is possible to gain considerable

information, at least of a comparative kind, from such records as

I had at my disposal.

Uncertainty of age and varying conditions of health on arrival,

and the abnormal conditions during detention, make it doubtful

if the maximum durations at the Gardens give very reliable

evidence as to the potential longevities of the species concerned.

The following conclusions, however, have some foundation, at

least if taken relatively. Amongst Primates, the Anthropoids

have a lower potential longevity than man, and _ longevity

decreases gradually towards the smallest monkeys, whilst in the

case of Lemurs it rises considerably. ‘The potential longevity of

Carnivora is relatively high, and wherever the conditions are

reasonably favourable, cases of long lite in proportion to size are

abundant. Insectivora have low potential longevity. Rodents

have an unexpectedly high potential longevity, there being many

cases of unusually great age, in proportion to size. Elephants

and the larger Ungulates have low potential longevities in pro-

portion to their size; all Ruminants have a relatively low

potential longevity, and the correlation with size is absent, or

almost reversed; in proportion to their size, many of the smaller

forms, such as Sheep and Goats, are much longer lhved than

Giraffes, Elands, or Cattle. Hdentates supply a number of

examples pointing toa high potential longevity; whilst Marsupials

generally, with exceptions, such as Wombats and Phalangers,

have a relatively low potential longevity.

Metchnikoff has elaborated the view that a chief cause of

senility is auto-intoxication from intestinal putrefaction, and

that, on the whole, those animals with the most capacious ceca,

large intestines and rectums, have the shortest lives in proportion

to their size and organization. The inferences from the maximum

and average durations recorded here, are in general agreement

with this view; the number of instances of animals and groups

of animals with relatively capacious hind-guts and short and

fragile lives being very great.. The most notable exceptions are

some of the Rodents, for instance Squirrels and Porcupines, and

some of the Marsupials, such as Wombats and Phalangers, which,

although they have relatively capacious hind-guts, have relatively

high viability and longevity. It must be noted, however, that if

the data were such as to exclude the unfavourable effect of the

special conditions in captivity, these exceptions might disappear.

I think the general proposition is true, that in proportion to

its size, an animal with a short hind-gut is hardier and has an

expectation of longer life.

The most important inference that I draw from these records

is the complete failure in practice of the theory that protection

from cold by artificial heat is the fundamental requisite for

mammals in captivity. - In the long list of mammals, belonging to

nearly every group, and coming from every part of the world,

there is no case of the successful application of this method, that

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 469

is to say, there is no case where the maximum duration is near

what might be expected, and the average duration bears a

reasonable relation to the maximum. Individual exceptions, or

knowledge from other sources, show us the ages to which animals

may attain; the records show the complete failure to get near

what is possible. Where comparative experiments have been

made the contrast is amazing. Compare the large cats, which

were relatively well housed, with the small cats, which were

confined in an overheated house without access to open air;

compare the civets, which were kept out of doors, with the genets

and paradoxures, kept in a heated house; the larger canines with

the smaller forms; the ratels and badgers (unheated) with other

mustelines (heated); the raccoons (unheated) with coatis (heated);

the squirrels and porcupines (unheated) with most of the other

rodents (heated); the larger marsupials and wombats (unheated)

with the smaller marsupials (heated).

The most fatal type of housing for any mammal (or bird) is

being confined to the interior of a warmed house, without free

access to the open air. The conditions of course are very

complex, and I do not suggest that the provision of heat is in

itself an evil. The ideal throughout the period I am discussing,

and which still maintains an evil existence in the minds of a

majority of those who have to do with living animals, is that, in

the first place, animals have to be kept warm. Warmth having

been secured, the more advanced persons have consented to, or

even urged, the advantage of ventilation, moistening of the air,

and so forth. The idea, however, is wrong. The first requisite

is free access to the open air, the next is light, space and clean-

liness ; these things having been secured, any form of heating

that may be thought advisable, may be added i so far as it does

not in any way interfere with the primary considerations. So far

as I can judge from my experience and observations, the main-

tenance of an equable temperature, and probably even a greater

amount of heat than is usually provided, is important in the case

of reptiles and of young or sick mammals or birds. Adult

mammals and birds, if in a normal state of health and if provided

with tolerable space for exercise, not only do not require an

equable temperature, but thrive better if they are subjected to

changes of temperature. The best possible conditions for them

are free access to large open-air enclosures, with adequate wind-

screens, and with the possibility of retreat to small, dry shelters,

which may or may not be provided with some form of artificial

heat. The smaller such shelters are the better, and certainly they

should not be, whether heated or unheated, part of a building to

which visitors have access. In a much larger number of cases

than is generally supposed, if such shelters are small and dry, and

provided with appropriate bedding, artificial heat is unnecessary.

Mammals and birds, if they have room for exercise in fresh air,

not only can maintain the heat of their bodies, but as a direct

reaction to the stimulation of air and change of temperature,

Proc. Zoou, Soc.—1911, No. XXXI. 31

470 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

improve in general vitality and in the quality of their hair, fur,

or feathers. Their viability is in a large measure determined by

their power of resistance to bacterial and probably other parasites.

It is impossible to ward off infection altogether, but the general

condition of the body and exposure to light and air are the most

certain preservatives.

It appears to be certain that although infection by a very few

parasites (microscopic or macroscopic), as these are living

creatures with the power of reproduction, is theoretically

enough to contaminate an animal, yet, in actual practice, a fairly

heavy bombardment ef parasites is necessary. Confinement in

heated houses not only weakens the natural power of resistance,

but affords the greatest opportunity for heavy bombardment.

Exposure to cold, light and air rapidly destroy a vast proportion

of the parasites or their eggs or embryoes that are not in actual

occupation of their host, and warmed houses afford the best

possible environment for them. I do not doubt, that by reducing

artificial heating as much as possible, and by making the shelters

as small and as easy to clean as possible, the damaging effect of

parasitic infection could be reduced enormously.

AVES.

PASSERES.

TuRDIDS. iwiy, ae Toe

LRANS, WALOIS REMOS oncsagoosossobsooones 68 46°7 154

T. albwentris (White-bellied Thrush). 2 10°5 20

fl, cemralos (Giseyy WweHS!0)) sotaccdcotesoos 2 52°5 64

T. falklandicus (Falkland Island 1 32 32

Thrush).

T. flavipes (Brazilian Grey Ousel) 1 30 30

Lh, UsaReons. (IESG MINE) so daesodeacesdso0n 5. 9 40 75

T. leucomelas (Brown Thrush) ......... 5) 67 119

St, coerulea (TElevels arte!) eokedadooconnesbac 7 D9 135

T. migratorius (American Robin)...... Ht 61 154

T. musicus (Song-Thrush)............... 4 74 ayy

T. nvustelinus (Wood-Thrush) ......... 4 14 29

1’. pocilopterus (Grey-winged Blackbird). 1 12 12

SN spUlkaarOS) ((TMEIGHENE®)) noosesnoooseoo05055%0 8 29 53

T. rufiventris (Red-bellied Thrush)... 3 67 95

T. torquatus (Ring-Ousel) ............... 5 47 93

AN, rarosies (storey AUIGVAUISIN)) ~ Soonccokoas458 2 57 103

T. viscivorus (Missel Thrush) ......... 3 8 12

Geocichlacitrina(Orange-headedGround 2 27 50

Thrush),

G. wardi (Pied Ground Thrush) ...... i 12 12

Monticola cyanus (Solitary Thrush)... 7 18 46

M. saxatilis (Rock As BRN, LO) 24°5 40

Myiophonus horsfieldi (Horsfield’s 1 62 62

Whistling Thrush).

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

No. of Max. dur.

471

Av. dur

TuRDID& (cont.). Indiv. in months. in months

Saaicola enanthe (Wheatear) ......... 11 12 62

Pratincola rubetra (Whinchat)......... 7 9 22

Ruticilla phenicurus.(Redstart) ...... A 4 ia

R. tithys (Black Redstart)............... 1 4 4

Cyanecula suecica (Blue-throat) ...... 1 27 27

Erithacus rubecula (Red-breast) ...... 4 10 16

Miro albifrons (South Island Robin). 4 9 28

Copsychus saularis (Indian Dial-Bird). 9 39 133

Cittocincla macrura (Indian Shama)... 4 46 53

Sialia wilsont (Common Bluebird) ... 29 20 54

Daulias luscinia (Nightingale)......... 21 16 AD

Sylora, whole Genus) ..... 5.26.26. cece eee 21 9 27

S. atricapilla (Blackeap) ....... Perea 13 10 27

S. cinerea (Whitethroat) ............. A Meb sich 9 20

S. currwea (Lesser Whitethroat) ...... 3 8 14

S. sylvicola (Wood Warbler) ............ 1 5 5

Secrochelus$Wallow, % 5), $iesen2s---- 1 2 2

Acrocephalus streperus(Reed-Warbler). 1 5 5

Accentor collaris (Alpine Accentor) ... 11 23 52

A, modularis (Wood Sparrow) ......... 1 4] Al

Mimus polyglottus (Northern Mocking 26 23 79

Bird).

M. saturninus (Saturnine ntgcehan 3 31 AT

Bird).

Galeoscoptes carolinensis (Cat-bird) ,.. 3 19 51

Harporhynchus rufus (Brown Mock 3 31 71

Thrush).

Cichlherminia densirostris (White-eyed 1 16 16

Mock Thrush).

CINCLIDA,

Cinclus aquaticus (Dipper) ............ 7 7 26

PANURIDE.

Panurus biarmicus (Bearded Tit-. 14 8 31

mouse).

PARIDE.

Parus, whole genus .......... ns EE RO 5D 18 108

Paaier (Coal Ditmouse)\, 0.255.922. 4 16 30

P. borealis (Northern Titmouse)...... 5 32 73

P. caudatus (Long-tailed ,, _ )...... 2 4 5

P. ceruleus (Blue RARE) O. ss. 9 18 29

P. cristatas (Crested Es aint 2 1:5 2

P. cyanus (Azure 9 ees: 3 11 12

P. major (Great 3 yet oe 14 21 108

P. palustris (Marsh i stds: 3 18 AO)

P. varius (Red-sided ,, | sles is: 15 36

Liothrix luteus (Yellow-bellied 4] 27 100

Liothrix).

ok*

472 DR. P. CHALMERS MITCHELL ON LONGEVITY AND j

SITTID a! a au ene

Sitta cesia (Common Nuthatch) ...... 11 10 d7

MOorACILLIDA.

Motacilla, whole genus ...............-.. 28 yy 60

AT. flava (Blue-headed Wagtail) ...... G 11 27

M. lugubris (Pied Sang) aide ieee 10 13 45

M. melanope (Grey Bede) teh Shek 8 24 60

M. raii (Yellow Py ib tenet 3 a8 51

Anthus, whole genus ..........-.-.---.--- 10 13°8 43

rAarboredsn nee seapit) sian ska ee 4 14 24

Ancobscurgis (UROCK) Ie.0))-cdcnasia nace ee 2 28-5 43

A. pratensis (Meadow Pipit) ............ 3 7 18

A, spipoletta (Water ANN aA aa sees 1 2 2 |

Trallina australis (Australian Pipit) . 2 il 20

PYCNONOTIDA. |

Pycnonotus, whole genus .............-- 4} 31 132

P. atricapillus (Black-capped Bulbul). 1 12 12

P. crocorrhous (Yellow-vented ,, ). 1 15 15

P. hemorrhous (Red-vented op 46 132

P. jocosus (Red-eared 5, dee LO 26°6 79

P. leucotis (Wihite-eared ) 55> ))). 98 Some 91

P. sinensis (Syrian Nes ae Nit 23

P. xanthopygos (Chinese Hae) es 24 83

Hemixos favala (Brown-eared ,, ). 3 27 62

Hypsipetes maclellandi (Rufous-bellied 2 un 114

Bulbul).

Chloropsis aurifrons (Malabar Green 3 22 39

Bulbul).

C. hardwickii (Blue-winged Bulbul)... 1 130 130

CRATEROPODIDA.

Garrulax, whole genus .........-........ 26 50 107

G. albogularis (White-throated Jay- 3 15 a

Thrush).

G. chinensis (Chinese Jay-Thrush) ... 11 61 107

G'. lewcolophus (White-crested Jay- 7 AT 98

Thrush).

G. pectoralis (Black-gorgetted Jay- 2 3 42

; Thrush).

G. perspicillatus (Masked Jay-Thrush). 2 91 103

G. picticollis (Collared *, 1 97 97

Turnagra crassirostris (South Island 5 15-4 26

Thrush).

Schee capistrata (Black-headed Sibia). 3 44 61

Leucodioptron canorum (Melodious 6 30 81

Jay-Thrush).

RELATIVE VIABILITY IN MAMMALS AND BIRDs.

CRATEROPODID (cont.).

Grammatoptila striata (Striated Jay-

Thrush).

Struthidea cinerea (Grey Struthidea).

Hypocolius ampelinus (Grey Hypo-

colius).

ORIOLID&.

Sericulus melinus (Regent Bird) ......

Oriolus, whole genus ................0000

O. indicus (Black-naped Oriole) ......

O. kundoo (Sykes’s Vitesse

O. melanocephalus ( Black- headed Or iole)

DIcRURID.

Chibia hottentotta (Indian Drongo) ...

Artamus, whole genus ..................

A. fuscus (Dull Wood Swallow) ......

A. personata (Masked Wood Swallow).

A. sordidus (Sordid ¥

A. superciliosus (White- -eyebr owed

Wood Swallow).

LANIIDE.

Hanius, whole genus ........4......-.+-.-

L. collurio (Red-backedShrike) ......

LL. eacubitor (Great Grey ,, ) ......

L.lahtora (Indian ,, saya

Laniarius quadricolor (¥ our-coloured

Shrike).

Cochoa viridis (Green Cochoa) .........

AMPELIDE.

Ampelis cedrorum (Cedar Bird) ......

A. garrulus (Waxwing) ..............0655

MUscICcAPID.

Muscicapa atricapilla (Pied Flycatcher).

HIRUNDINIDE.

Hirundo rustica (Swallow)...............

NECTARINIID.

Zosterops, whole venus ...............64

Z. dorsalis (Grey-backed White-eye) .

Z. lateralis (Lateral 4 )

Z. palpebrosus (Indian saab it 2

Z. simplex (Chinese aay ey)

No. of

Indiv.

HH Ot

re os ee Or Or

Av. dur.

in months.

473

Max. dur.

in months.

ATA DR. P. CHALMERS MITCHELL ON LONGEVITY AND _

No..of Ay. dur. Max. dur.

MELIPHAGIDS. Indiv. in months. in months.

Prosthemadera nove-zealandie (Poé 19 14 58

Honey-eater).

Myzantha garrula (Garrulous ,, ). 1 9 9

Anthornis melanura (Black-tailed 5 6°4 13

Flower-bird).

Xanthomyza phrygia (Warty-faced 5 Ug) 37

Honey- eater).

Entomyza cyanotis(Blue-faced Honey- 4 5 12

eater).

C@REBIDE.

Careba cyanea (Yellow-winged Sugar- 26 24:4 92

bird).

Dacnis cyana (Blue Sugar-bird) ...... 4 11:5 Ie

Chlorophonia spiza (Black-headed 2 42°5 84

Sugar-bird).

TANAGRIDAS.

Chlorophoniu viridis (All-green Tanager). 6 3 4

Huphonia, whole genus ................+ 4] 12°5 39

EH. chlorotica (Greenish Tanager) ...... 1 1 1

L. flavifrons (Yellow-fronted Tanager). 2 16 30

Lf. lanvirostris (Thick-billed Boe ee ease) gt 39

H. nigricollis (Black-necked ,, ). 4 (a) 10

EL. pectoralis (Pectoral ye) ak cea 8 8

E. sclatert (Sclater’s Si tha eek | 1 if

E. violacea (Violet Sh hae 15 3

Hypophea chalybea (Lead-coloured 1 11 11

Tanager).

Tanagrella cyanomelena (Blue & Black 2 145 Sd

Tanager).

T. velia (Red-bellied Tanager) ......... 1 2 2

Calliste, whole genus .................0.0+ 32 115 39

C. brasiliensis (Brazilian Tanager) . 3 4) 8

C. fastuosa (Superb Tanager) ...... 13 15 23)

C. festiva (ReStivensitae nme) uy eee 3 4 4

C’. melanota (Black-shouldered Tanager). 2 5 6

C. tricolor (Green-headed Tanager)... 11 11°8 3:

Tanagra, whole genus .................. 24 16 70

T. cana (Silver-blue Tanager) ......... 3 12 23

T. cyanoptera (Blue-shouldered Tanager) 7 - 18 49

T. palmarum (Palm Tanager) .......-. 5 11 33

T. sayaca (SEROUS) Soon adcca 2 7 il

UE si iaianin (Surlabe dere) )) ener ff 21 70

Stephanophorus leucocephalus (White- 2 3 4

capped Tanager)

Tachyphonus coronatus (Crowned 1 8 8

Tanager).

T. melaleucus (Black Tanager) ......... 8: 22, 47

RELATIVE VIABILITY IN MAMMALS AND BikDS.

TANAGRID (cont.). ee

Pyranga saira (Saira Tanager) ......... 2

Rhamphocelus brasilius (Brazilian 13

Tanager).

Cissopis leveriana (Magpie Tanager) 9

Saltator auranturostris (Ovange-bellied — 1

Tanager).

S. magnus (Great Tanager) ............. 1

ersiieies \(CAUMERG 550) anteater 2

Orchesticus ater (Black-headed Tanager). 1

PLOCEIDA.

fistrelda, whole genus...........1¢.0...04- 176

£. amandava (Amaduvade Finch)...... 30

#. bella (Beautiful Finch) ............... 6

E. bichenovit (Bicheno’s Finch)......... 8

E. cinerea (Common Waxbill)...... 21

#. cerulescens(Cinereous ,, ) ...... 5

EH .cyanogastra(Blue-breasted Waxbill) 6

EL. dufresnit (Dufresne’s a otal

H. formosa (Green Eada) ae 6

#. granatinag (Grenadier Ss eee

E. melpoda (Orange-cheeked ., ) 17

. phenicotis (Crimson-eared ,, ) 8

EH. phaéton (Crimson nS MR

#. rubriventris (Red-bellied ee aay. al

EL. squamifrons (Scaly-fronted - ,, ) 4

Hi. subflava = (ebra sega ae

E. temporalis (Australian Hera) #310)

Spermestes, whole genus ............... 26

S. cucullata (Hooded Finch) ............ 10

S. fringilloides (Pied Grass-Finch) ... 12

Se qanven (Wwart Winch) a/c.) 2 h8)..2: 4

Pytelia, whole Semus! 22.) ioe68 5. caee 9

P. citerior (Western Melba Waxbill)... 1

P. phenicoptera (Crimson-winged 6

Waxbill).

P. wienert (Wiener’s Finch) ............

Amadina, whole genus .........0..225.6 €

A. bicolor (Cape Palmas Finch).........

A. castanotis (Chestnut-eared Finch)

A. erythrocephala (Red-headed

)

A, fasciata (Cut-throat Bo 8)

A. fringilloides (Fringillid ay)

A. lathami (Spotted-sided 5, -)

A. modesta (Modest sakes)

Munia, whole-wenus .2:.....020.08 0040: )

M. ferruginea (Javan Maja Finch) ...

~H! 1 :

met OUD OF-! OV AT =1 U5 OO bO

Ay. dur.

in months.

475

Max. dur.

in months.

476 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

PLOCEID& (cont.).

Munia maja (Maja Finch)

M. malabarica (Indian Silver-bill) ...

M. malacca (Black-headed Finch) ......

M. nisoria (Bar-breasted ieee

M. punctularia (Nutmeg-Bird) .........

M. rubro-nigra (Chestnut-bellied

Finch).

M. striata (Striated Finch) ............

Macopelas (Mopela, | 9h) aeeneeenere

Pocphila, whole @enus..-...-.-..4:42-020-

P.acuticauda (Long-tailed Grass- Finch)

P. cincta (Banded a5 any)

P. gouldie (Gouldian “4 se)

P. mirabilis (Beautiful a Fe

Donacola castaneothorax (Chestnut-

breasted Finch).

D. pectoralis (White-breasted bat i)

Padda oryzivora (Java Sparrow) ......

Erythrura prasina (Fire-tailed Finch)

EL. psittacea (Parrot se)

Vadtia wioleycemis sen. aeeeeet eee. cee eee

V. albonotata (White-marked Whydah

Bird).

V. ardens (Red-chested Whydah Bird)

V. paradisea (Paradise _,, 6

V. principalis (Pin-tailed _,, )

Chera progne (Long-tailed Weaver-

bird).

Hypochera nitens (Shining Weaver-bird )

Coliopasser macrurus (Yellow-backed

Whydah Bird).

Urobrachya, whole genus ...............

U.albonotata (White-winged Whydah

Bird).

U. axillaris (Red-shouldered Weaver-

bird}.

U. bocagit (Orange-shouldered Weaver-

bird).

Huplectes, whole genus ..............04..

FE. afer (Black-bellied Weaver-bird).

LE. capensis (Y ellow-shouldered Weaver-

bird).

E. flammiceps (Crimson-crowned

‘Weaver-bird).

EL. nigriventris (Black-chested _ ,, )

E. oryx (Grenadier

Pyromelana aurea (Golden- backed

Weaver-bird).

No. of

Indiv.

Av. dur.

in mouths.

Max. dur.

in months.

101

115

109

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

PLocEID& (cont ).

Pyromelana franciscana (Scarlet

Weaver-bird.)

Poudiaerythrops (Red-faced ,,

ff. madagascariensis (Red- “headed

Weaver-bird).

Pyrenestes albifrons (White-fronted

Weaver-bird).

Spermospiza hematina (Blue-beaked

Weaver-bird).

S. guttata (Bright-spotted _,, bala)

Quelea, whole genus .....2.¢.22cce000e40s

Q. occidentalis (Black-cheeked Weaver-

bird.)

Q. russi (Russ’s | Weaver-bird)

Q. sanguintrostris (Red-beaked _,, )

Floceus, whole genus’ ..).........8..0:-.25

P. atrigula (Black-throated Weaver-

bird).

P. bengalensis (Bengal Weaver-bird)

P.manyar ° (Manyar _,,

Heterhyphantes melanogaster (Black-

chested Weaver-bird).

Sitagra luteola (Yellowish _,, oe)

ITyphantornis, whole genus ............

H. brachyptera (Short-winged Weaver-

bird).

Hf. capensis (Olive Weaver-bird) ......

HT. castaneofuscus (Chestnut-backed

W eaver-bird).

HH. nigriceps (Black-headed _ ,, ae)

H. personata (Masked lag sale)

H. superciliosus (Kye-browed ,, ,, )

H. textor (Rufous-necked ,, ,, )

H. velatus (Black-fronted ,, ,, )

Lenton alectory(Ox= id) 25.25.24 eee

FRINGILLIDZ.

Cyanospiza cris (Nonpareil Finch) ...

CNeyonea Gndiszo-bird)\y cccertonadten<e:

Phonipara canora (Melodious Finch)...

P. olivacea (Olive Finch) ...............

Paroaria, whole genus ................-.

P. capitata (Yellow-billed Cardinal) ..

P. cucullata (Red-crested pins ae

P. larvata (Red-headed ss

Gubernatria cristatellus (Black- “crested

Cardinal).

Diuca grisea (Diuca Finch) ............

No. of

Indiv.

oD)

—_

me OT OLD Os <O

—

bo (se)

MeN OUD ON ON tS

—

Avy. dur.

in months.

477

Max. dur.

in months.

102

A478 DR. P. CHALMERS MITCHELL ON LONGHVITY AND

FRINGILLIDE (cont. ). ae see ae

Phrygilus, whole genus .................. 4 31 86

P. alawdinus (Alaudine Finch)......... 1 86 86

IP. Frutice) pa Orchercd Gin yaw )ene cee 1 15 15

P. gayt (Gay’s 55) isndieetasicie 2 27 37

Guiraca, whole genus .............++2++++ 12 21 v1

G. cerulea (Blue Grosbeak) ............ 2 26 35

G. cyanea (Brazilian Blue Chi osbeak)... 8 Lg) al

G. parellina (Lazuline Finch) ......... 2 25 4D

Hedymeles ludovicianus (Rose-breasted — 2 0 11

Grosbeak).

Pheucticus chrysogaster (Yellow-bellied 1 oi Bil

Grosbeak).

Cardinalis virginianus (Car dinalig. ) )) 22 31 83

Oryzoborus crassirostris (Thick-billed 1 53 D3

Seed-Finch).

O. torridus (‘Tropical Hs 7 21 4

Coryphospingws cristatus (Red-creasted 8 AT 123

Finch).

O. pileatus (Pileated Finch)......... 5 14 AO

Spermophila, whole genus ............... 50 43°5 WAT

S. albogularis (White-throated Finch) 5 83 123

S. cerulescens (Bluish shee aa 43 78

S. collaria (Collared Pee ar 2 109 127

iS. eulert (Euler’s on) RN 2 2

S. gutturalis (Guttural Ly Dwele 30°5 34

S. hypolewca (Half-white m 2 16°5 33

S. lineata (Lineated iva) nee 66 88

S. lineola (Lined Jil Ep toll 36 110

S. nigro-aurantia (Reddish eh) Veo 45 101

S. plumbea (Plumbeous jen): tei a6 85

S. torqueola (Black-banded a § re 3 3

Basser wivolevee musi yiber sede. eee sack 25 22 135

P. alario (Alario Sparrow) ............ 12 8 16

JB, ommeunauis (CRO gy) so aeaasanace 1 21 21

P. diffusus (Black-breasted Sparrow)... 1 19 19

P. luteus (Yellow BN) get 8) AO 124

P. montanus (Tree parities) oh Hee 16 ay)

P. salicicola (Spanish SMa) 1 38 3

Jeg simplex (Grey-headed _,, 3 45 135

Petronia petronella (Yellow-thr oated 7 Se Thigh

Rock-Sparrow).

P. siulta (Rock-Sparrow) ....)..:.-.---- 2 A6 91

Coccothraustes, whole genus ............ 31 22°5 92

C. melanurus (Black-tailed Hawfinch) 8 34 74

C. personatus (Masked sah yee 12°5 24

C. vulgaris (Hawiineh) ,.........-00..+5+: 21 Ig) ue.

Mycerobas melanowanthus (Black enol | Il Ad 44

Yellow Hawfinch). Shas

Raat inus chloris (Greenfinch) 2 38 66

RELATIVE VIABILITY IN MAMMALS AND BIRLS. 479

No. of Av. dur. Max. dur.

FRINGILLIDE (cont.). Indiv. in months. in months.

Ligurinus sinicus (Japanese Green- 5 | 27 85

finch).

Montifringilla nivalis (Snow-Finch)... 2 21°5 37

Tingula, whole genus .....s.se0ccee0e03 43 36 142

eromleds: (Chaitin) oh... asceson<- cok! 7 33 142

FE. kawarahiba (Kawarahiba Finch)... 2 42 5D

F. Vinaria (Mealy Redpoll) ............ 1 89 89

Ff. montifringilla (Brambling) ......... 22 AD 105

FI’. spodiogenia (Alpine Chaffinch)...... 2 23 24

FF. teydea (Reydeant) 770) -ess: 5 10 22

Beiiiilon (VTintillon’ 4, .\)...2s 4 23 60

Carduelis elegans (Goldfinch)............ 21 24 57

Corientans(Kastern ,, )..........+. 3 18 43

Chrysomitris, whole genus ............... 33 13 90

C. barbatus (Black-chinned Siskin) ... 5 14 4

C. spinoides (Indian 4 REN A 90 90

Cmepinus, (SISKIM) +... eosherec seat 23 eZ 74

C. tristis (American Siskin) ............ 3 1] 32

Cs yarreia, (VY amrells) BY ye anes 1 24 24

Sycalis, whole genus. ............-0.-0005- 32 27 90

S. arvensis (Field Saffron Finch) ...... 2 33 52

S. flaveola Cais Bp) eek t- 22 24 90

S. luteola (Yellowash~ (ie) Rtas. 8 dl 80

Crithagra, whole genus ...............+++ 34 24 90

C. albogularis (White-throated Seed- 6 10 30

eater).

C. butyracea (Yellow Seed-eater) 10 22 90

C. chrysopyga (Yellow-rumped ,, ) 13 38 80

C. musicus (Singing ai 5) mad Li, 24

C. sulphurata (Sulphury Oy Eyl )eteais 2 4

Serinus, Whole genus ..........0s.02-08+- 18 15 52

S. angolensis (Angolan Serin) ......... 1 22 22

Sweananiisy (Canary ered. seteek cn =. 12 14 52

S. canicollis (Grey-necked Serin) ...... 1 20 20

IS hortulanis (Serin) .05..25..40 see. ns 3 10°3 24

S. tottus (Brown Canary) .........2..... 1 27 27

Linota, whole genus ...............00.65- 28 ab 92

L. cannabina (Linnet) ........2....0002. 3 14 24

Ls flaninosirisl (Witte) Py. c.1e6 ees. cnesks 4 39 92

L. rufescens (Lesser Redpoll)............ 21 18°5 74

Carpodacus erythrinus (Ruddy Finch) 13 28 79

C. hemorrhous (Blood-stained Te) anne! 25 49

Erythrospizagithaginea(Rosy Bullfinch) 6 2 52

Pyrrhula europea (Bullfinch)............ 8 12 4]

P. rubicilla (Siberian Bullfinch) ...... 7 12 23

Pinicola enucleator (Pine-Grosbeak)... 10 13 44

Bovia, whole Cenuss, 9, 5A: eee... 29 8 39

L. pityopsiitacus (Parrot Crossbill) ... 3 24 39

L, leucoptera (White-winged Crosshill) 4 12 37

480 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Av. dur. Max. dur.

FRINGILLIDE (cont.). Indiv. in months. in months.

Loxia curvirostra (Common Crossbill). 21 6 24

L. bifasciata (Kuropean White-winged 1 5 5

Crossbill).

Plectrophenax nivalis (Snow Bunting) 63 11:5 65

Calcarius lapponicus (Lapland ,, ) 4 18 30

Fimberiza, whole genus .................. 86 36 165

L. aureola (Yellow-breasted Bunting) 4 85:5 115

Li. cia (Meadow sox) i een 7 i

E. ciopsis (Bonaparte’s ray eat a ist 11

E. cirlus (Cirl ee) all 21 63

L. citrinella (Yellow Hammer)......... 4) 27 Hath

L.. chrysophrys(Yellow-browed Bunting) 2 D3 86

EL. hortulana (Ortolan Dee |) erst) 25 15

L. luteola (Red-headed ,, ) 11 63 165

E. melanocephala (Black-headed ,, ) 7 52 141

LH. miliaria (Corn agitate 45 123

EL. rutila (Red-backed ,, ) 5 50 149

H. scheniclus (Reed 7 eG 20 62

H. striolata (Striolated Pow ee 15 21

Huspiza americana (American ,, ) 3 15 23

fringillaria sahare (Sahara ta hee 25°5 38

Melophus melanicterus (Crested Black 2 12°5 24

Bunting).

Zonotrichia albicollis (White-throated 2 W3 on

Song Sparrow).

4. pileata (Pileated _,, sae ye 17 42

Junco hyemalis (Snow-bird) ...........: if ii 72

Spizella socialis (Chipping Sparrow)... 1 8 8

Pipilo erythrophthalmus (Red-eyed 4 25 A5

Ground-Finch).

ICTERID#.

Ostinops decwmanus (Crested 2 21 28

Hangnest).

Cassicus hematorrhous (Red-rumped 2 Al 57

Hangnest).

C. persicus (Yellow Bs ye dl 46 102

Tetenus, wholeywenus /..24......00) 420-0. 31 27 110

L. abeillei (Black-sided Hangnest) 1 30 30

1. baltimore (Baltimore Se Sy Nees 14 38

I. chrysocephalus (Yellow-crowned 1 66 66

Tropical Hangnest),

1. jamaict (Brazilian Hangnest) ...... 7 40 80

L. spurius (Orchard x5 ee ohinte 6 17 31

L. tibialis (Yellow-shouldered Hang- 3 27 45

nest).

Z. vulgaris (Common Hangnest) ...... 8 27 110

Dolichonyx oryzivora (American Rice- 1 3 3

bird).

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 481

No.of Av. dur. Max. dur.

IcrERIDE (cont.). Indiv. in months. in months.

Molothrus, whole genus ...............0.. 25 65 145

M. badius (Bay Cow-bird) ...... 5 67 145

M. bonariensis (Silky eee) Pete 8: doe: 18 65 135

M. purpurascens (Purple ,, )~....... 2 52 71

Ageleus, whole genus ...........6...5+. 8 44 94

A. frontalis (Chestnut- -fronted Troupial). 1 16 16

A. pheniceus (Red-shouldered Starling) 5 54 94

A. ruficapillus (Red-headed Marsh- 2 33 65

bird

Xanthocephalus icterocephalus (vellow. 11 AT 123

headed Troupial).

Xanthosomus flavus (Yellow , ) 3 53 98

Amblyrhamphus holosericeus (Silky 7 50 122

Hangnest).

Pseudoleistes virescens (Dark-green 5 26 5D

Maize-eater).

Sturnella defilippi (De Filipprs 18 16 39

Meadow-Starling).

S. ludoviciana (Louisianian ,, = 2 10°5 16

Cureus aterrimus (Chilian Starling)... 1 24 24

Lampropsar dives (Rich-black i 2 2

Troupial).

Quiscalus lugubris (Black PA 1

Q. versicolor (Changeable ,, ) 9 10 3l

Aphobus chopi (Chopi Starling) ...... 62 102

STURNIDA.

Lamprocolius: auratus (Purple-headed 3 141 155

Glossy Starling).

L. chalybeus (Green _,, fp 5 71 150

Lamprotornis cneus (Long-tailed 6 44 17

Glossy Starling).

Sturnus menzbiert (Indian Starling) 2 25 26

S. vulgaris (Common aan) 9 26 58

Sturnopastor contra (Pied Mynah) ... 3 D9 107

Acridotheres, whole genus ............... 25 44 145

A. cristatellus (Chinese Mynah) ...... 6 o4 62

A. fuscus (Brown stabi isis eu 6 42 145

A. ginginianus (Indian ,, ) ...... 1 4] 4]

A. mahrattensis (Southern Brown 2 58 97

Mynah).

A. tristis (Common Mynah) ............ 10 47°8 117

Poliopsar, whole genus .................- 8 30 a6

P. andananensis (Andaman Starling). 2 20°5 38

P. blythi (Blyth’s 1 23 23

P. malabaricus (Malabar Mynah) fe 48) 33 97

Temenuchus pagodarum (Black-headed = 1 13 ils:

Mynah).

482 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Av. dur. Max. dur.

STURNIDH (cont. ). Indiv. in months. in months.

Pastor roseus (Rose-coloured Pastor) . 24 34 on

Gracupica nigricollis (Black- anokel — 1 16 16

Grackle).

Dilophus carunculatus (Wattled 6 42 73

Starling).

Gracula, whole genus ..... MEATS Poe 39 30 124

G. intermedia (Larger Hill Mynah) ... 20 33 124

G. javanensis (Javan Mynah) ......... 1 77 ei

G. religiosa (Small Hill Mynah) ...... 18 22 61

PTILOCNORHYNCHIDZ.

Piilonorhynchus smithti (Green Bower 1 19 19

Bird).

P. violaceus (Silky Bower Bird) ..... ep BO) BO): 115

Chlamydodera maculata (Spotted 5 AQ 68

Bower Bird).

PARADISEIDE.

Paradisea minor (Lesser Bird of 5 30 65

Paradise).

P. rubra (Red Bird of Paradise) ...... i 4 A

Ptilorhis paradisea (Rifle-bird) ...... 1 46 46

Seleucides nigricans (Twelve-wired 1 11 11

Bird of Paradise).

Manucodia chalybea (Green Manucode) 1 12 12

CoRvIDE.

Corvus, whole{genus ..0....ac.6c.20002 5. 140 24°6 133

CO. americanus (American Crow) ...... 1 5] 51

O. australis ‘(Australian ,, )....... 7 13 82

O. capellanus (Chaplain ,, ) ...... 23 22 89

OF coraw (Raven) oi Rr nie SUERTE 31 itil AZ

C. cornix (Hooded Crow)" eee. 14 35 99

OL corona (OREN 4, )) San6Bbnbodooece 12 1s 46

O. culminatus (Long-billed Crow) ... 2 20 36

OQ. daiiricus (Black-and-white Jackdaw).. 1 20 20

Orugilegus (UOols) reat sesne- eer eee 1S 10 QE

C. monedula (Jackdaw) .................. 15 39 89

O. scapulatus (White-necked Crow) .... 10 Dilsroisy wculilees

C. splendens (Indian Weoy el 38 133

Corvultur albicollis (White- Epoled il 18 18

Raven).

Nucifraga caryocatactes (Nutcracker). 6 27 93

Pica witole @enus oy s-cee.. sae ses see 19 51 131

Be hooters (Himalayan Magpie) .. 1 6 6

P. mauritanica (Moorish 7 eee Q: 9

Pr gaistica (Niappie)) ies. -eaee ean 15 54 13H

2 69 5)

P. sericea (Chinese Mag gpie) . / eee

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

Corvip& (cont.).

Cyanopolius cooki (Spanish Blue

Magpie).

C. eyanus (Chinese Blue Magpie)

Garrulus glandarius (Jay) ............065

G'. lanceolatus (Lanceolated Jay)

Cyanocitta, whole genus..................

CO. coronata (Crowned Jay)

C. cristata (Blue

99 J seme nen nnvne

CO. melanocyanea (Guatemala Blue Jay).

C. yucatanica (Yucatan

Cyanocorax, whole genus ...............

C. cyanomelas (Black-headed Jay)

C. luxuosus (Mexican =)

C’. cyanopogon (Blue-bearded ,, )

C. pileatus —_(Pileated a)

Calocitta formosa (Swainson’s Long-

tailed Jay),

9? 9?

Urocissa, whole SenUs 9 2...5..2..6.06. 00s

U. flavirostris (Yellow-billed Blue Pie)

U. magnirostris (Siamese ui) 73x.)

U. occipitalis (Occipital ape)

U. sinensis (Chinese vane

Cissa venatoria (Hunting Crow)

Dendrocitta, whole genus ...............

D. himalayensis (Himalayan Tree Pie)

D. sinensis (Chinese aay ss

D. vagabunda (Wandering ,,_ ,, )

Ptilostomus senegalensis (Piapec) ......

Pyrrhocorax alpinus (Alpine Chough).

Re gracules (howe ly passes: sree.

Corcorax melanorhamphus (White-

winged Chough).

Heterolocha gouldi (Huia Bird)

Streperd, whole genus .....2.....2+600++

S. anaphonensis (Grey Crow-Shrike) .

S. graculina (Ried. Se) ie

S. fuliginosa (Sooty _,,

Gymnorhina leuconota (White- backed

Piping Crow).

eee cee

G. tibicen (Black-backed __,, $

Cracticus destructor (Long-billed

Butcher Crow).

C. picaius (Pied cs it)

ALAUDID.

Alauda, whole genus .............-..000+-

A. arborea (Wood-lark)

A. arvensis (Sky-lark)

No. of

Indiv.

—

PRKHonw%1cd@

— e a a for) —

Ae aw eorb SCmMonwmnwnrrranwmnnon.e woondr bs

to) er)

Av. dur.

in months.

Silvas

12°8

483

Max. dur.

in months.

ABA DR. P. CHALMERS MITCHELL ON LONGEVITY AND

E No. of — Av. dur. Max. dur.

ALAUDIDA (cont. ). Indiv. in months. in months.

Alauda coclivox(Sweet-voiced Lark)... 1 13 13

A. crassirostris (Yhick-billed ,, ) ... 1 24. 24

A. cristata (Crested - 7 15 86

A. gulgula (Indian Sky-lark) i 97 97

Calandrella beetica (Andalusian Short- 7 14 19

toed Lark).

O. brachydactyla (Short-toed Lark) ... 2 28 AT

Mirafra affinis (Meadow Bush-Lark). 1 14 14

M. cantillans (Pale Indian ,, St aha 195} Y

Melanocorypha, whole genus............ 20 11:5 81

M. calandra’ (Calandra Lark) ...... 10 14°6 81

M. mongolica (Chinese ws vu) Lee beet 2 17 32

M. yeltoniensis (Black soa. ese 8 6:5 14

Rhamphocorys clotbeyi (Clotbeys 3 13 19

Lark).

Pyrrhulauda verticalis (White-headed 6 16 35

Bullfinch Lark).

Otocorys alpestris (Shore Lark)......... 12 25°6 61

O. bilopha (Algerian ,, ae art 2 95 10

PITTIDA.

Pitta bengalensis (Bengal Pitta) ...... 1 22:5 24

P. strepitans (Noisy JAN ome 4 11 29

TYRANNIDE.

Machetornis vrivosa (Short-winged 2 45°5 76

: Tyrant).

Tenioptera nengeta (Pepoaza fe) i 5 5

Tyrannus melancholicus (Melancholy 1 67 67

Tyrant).

7. pipiri (American King-bird) ..... , 1 12 12

Pitangus sulphuratus (Sulphury 21 34 104

Tyrant).

Milvulus tyrannus (Fork-tailed ,, ). 1 38 38

CoriNnGIDé.

Chasmorhynchus niveus (Carunculated — 1 D3 53

Chatterer).

CO, nudicollis (Naked-throated Bell- 18 ATL 49

bird).

Cotinga cincta (Banded Cotinga) ...... 2 3l 7

Rupicola crocea (Cock of the Rock)... 8 8°D 0

DENDROCOLAPTIDA.

Furnarius rufus (Red Oven-bird). ... 2 aD 7

MENURIDE.

Menura superba (Liyre-bird) ............ 2 575 107

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 485

Summary of Passeres.

The first general question that requires consideration is the

value of the data on which this summary is based. Passerine

birds, except to those who pay special attention to them as pets,

are relatively little individualized, and in a large collection it

may well be that the history of one individual has been assigned

by mistake to another individual. I cannot doubt that such

mistakes have sometimes occurred, but I do not think that they

have had much effect on the figures. The bias they would give

would tend on the whole to lower the figures, for where there was

a doubt as to a dead individual in the Keeper’s mind, I find that

he was disposed to assume that it was the oldest example. More-

over, if the figures be run through in detail, it will be found that

very often high records occur in cases where the number of

examples was so few that confusion of individuals cannot have

taken place, whilst the period of 33 years covered is so long in

proportion to the numbers in most cases, that probably this

source of error is on the whole insignificant.

Mr. J. H. Gurney’s remarkable paper * has already made plain

that birds attain considerable ages, and the figures of maximum

duration that I am able to record are in no case improbable. It

is to be remembered that probably the vast majority of

Passerines are received at the Gardens in an adult condition,

that the age of an adult bird cannot be determined from inspec-

tion, and Semel ares the maximum durations recorded are in all

cases below, and may be much below the potential longevity. Mr.

Gurney gives the following as the oldest Passerines of which he

had been able to obtain information: Raven 69 years, Gymno-

rhina tibicen, 26 years, Nightingale, 25 years, Skylark 24, Gold-

finch 23, Grosbeak 21, Canary 20, Bullfinch 19, Weaver-bird 9;

and he sets down 20 to 24 years as the possible limit of most

Passerines in captivity. Amongst the Garden records, I may

select the following :—Grey Struthidea, nearly 19 years, Paradise

Whydah Bird, over 16 years, Diuca Finch, nearly 16 years,

Glossy Starlmg and American Robin, about 13 years, Bay Cow-

bird, over 12 years, an Indian Crow over 11 years, a Bulbul

11 years, a Great Titmouse 9 years, a Lyre-bird, nearly 9 years.

Mr. W. Chamberlain, F.Z.S., informs me that he bought an adult

Cardinal (Paroaria cucullata) in 1874; that in 1896 it showed

signs of age, chiefly difficulty with the autumn moult, and

thickening of the scales of the legs; that it died in 1899,

having lived in a cage for 25 years 4 months. It must be

noticed that if size be taken into consideration, the ages bear

no relation to the relative sizes of the birds which attained to

them, and moreover that Passeres as a whole, in comparison with

their size, live to much greater ages than is the case with

mammals. This is entirely in accordance with the views of

Metchnikoff, for Passerine birds in every case have the hind-gut

* **The Comparative Ages to which Birds live,” Ibis, 1899, p. 19.

Proc. Zoou. Soc.—1911, No. XXXII, 32

486 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

and cxca very small in proportion to the total length and capacity

of the alimentary canal. ‘They digest rapidly, void their excreta

at frequent intervals, and have very little opportunity for intes-

tinal putrefaction.

With regard to viability, the record of Passerines is on the

whole much better than in the case of mammals; that is to say,

the maximum duration is frequently high and is much more

nearly approached by the average duration. There are some

notable exceptions. First, it is most striking that common

British birds, like common British mammals, have very bad lives

in captivity. J am inclined to put this down to the same kind of

reasons. In the first place, it must frequently happen that birds

are damaged by capture, or that many weak examples are taken,

and these find their way quickly to the Gardens, there to die;

whereas in the case of birds captured in remote regions, the

deaths take place more frequently before arrival, and a selection

of the hardier individuals is thus made automatically. The case

would be more striking if I had included in the averages, the

considerable number of birds that died without living for a month

at the Gardens, a number relatively much greater in the case of

common European and British forms. It seems, moreover,

highly probable, that wild birds in this thickly populated country

have survived in proportion to their development of a repulsion

to the vicinity of man, a repulsion which only a small proportion

of them, and that only gradually, can overcome in captivity. It is

most remarkable how the average duration of such kinds as

English Thrushes, Whinchats, Redstarts, Wheatears, Warblers,

Tits, Nuthatches, Pipits, Shrikes, Linnets, Finches, Buntings,

is from 12 months to | or 2 months, whilst that of their nearest

foreign allies may be many times greater.

The viability, as shown by the average duration, has no

constant relation to size. On the whole, soft-billed birds, such as

the Tanagers, fed chiefly on fruit and prepared food, have been

less successt ul than Finches and Weavers, and insectivorous bir ds

have done less well than seed- and flesh-eaters s. Nor is there any

striking result to be obtained from consideration of the quarters

assigned, for the vast majority of these Passerine birds have

been kept under similar conditions. The Western Aviary, with

open-air flights and heated retreats, has been their headquarters,

although some of the small Finches, Tanagers, and Weaver Birds

have been in cages in the Parrot House. The evidence points to

Passerine birds having a good viability and a potential longevity,

even in the case of the smallest, reaching well over twenty years.

Although success has been greater than in the case of many

mammals, I think that this is to be expected from the constitu-

tions of these two sets of creatures, and that in proportion to

their viability, the success in the case of Passerines is still far

short of what might be achieved. Much has already been done

to improve the access to air in the case of all the Passeres.

Capt. Flower’s highest records are a Paradise Whydah Bird,

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 487

still alive after 9 years’ duration; an Amaduvade Finch after

7 years, a Black-headed Chestnut-Finch after 9 years, a Java

Sparrow of over 10 years’ duration, anda Red-crested Cardinal of

over 8 years’ duration. It is noteworthy that these figures are

comparable with my records and relate to species or genera which

have had long-lived examples here. Actually our records are

higher, but the number of individuals amongst which they

occurred is greater.

PICARIA.

\ be No. of Av. dur. Max. dur.

CAPRIMULGID&. Indiv. in months. in months.

Caprimulgus europeus (Nightjar)......... + 4 iG

I have omitted a number of cases that lived for less than a

month, but this is only another instance of failure with common

European birds.

PICIDE.

Dendrocopus major (Greater Spotted 16 9-8 33

Woodpecker).

Centurus tricolor (Banded B Det Seer 14 14

Melanerpes erythrocephalum (Red- 1 2 2

headed Woodpecker).

Leuconerpes candidus (White-headed 2 12 16

Woodpecker).

Gecinus viridis (Green Woodpecker)... 9 0 0

Colaptes auratus (Golden-winged 8 23 54

Woodpecker).

Tynx torquilla (Wryneck) ............... 5 0 0

Amongst the Woodpeckers and Wrynecks there is again

complete failure with British birds, whilst the other figures would

appear to show a low viability for the birds of this group,

appearing both in the average and maximum durations.

Oona No. of Av. dur. Max. dur.

: Indiv. in months. in months.

Colius, whole, genus 2.22. <c-00-sonsddsases 14 35 91

Ceupensis\(Caipe Coliy) 2 ease .ac5s-2 ae 10 34:4 91

C. castanotus (Chestnut-backed Coly)... 2 8:5 9

C. erythromelon(Red-cheeked ,, )... 1 66 66

C.nigricollis (Black-necked ,, )... 1 65 65

These small, chiefly frugivorous birds, have in proportion to their

size a good maximum and average duration, and must be inferred

to have a good viability. It is interesting to note that the whole

length of the intestines is remarkably short, although capacious,

and there can be little opportunity for intestinal putrefaction.

In the period under consideration the Colies were kept in the

Parrot House, without access to open air.

39%

488 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Av. dur. Max. dur.

ALCEDINIDE. Indiv. im months. in months.

Alcedo ispida (Kingfisher) ............... 27 4 24

Halcyon, whole gemus ..................... 10 23 A4

H. chloris (Green-headed Kingfisher) . 1 28 28

H. sanctus (Sacred ieee iG 32

H. smyrnensis (White-br easted u 26 26

Kingfisher)

H. vagans (New Zealand e a 30 A4

Dacelounnolersenustrreeessee cece eee 54 25°5 84

D. cervina (Buft Laughing Kingfisher). 2 28 34

D. gigantea ( 4 5 \eeaoll 25 84

D. leachi (Leach’s = De seg 58 58

Tn the usual fashion, the failure with the Common Kingfisher

has been very great. The other Kingfishers, in proportion to

their size, show poor maximum and average durations, and these

birds must be regarded as of relatively low viability. | Although

there are no ceca, the hind-gut is relatively long and capacious.

Mr. Gurney cites, from the Rotterdam Gardens, an instance of a

Laughing Kingfisher having lived for 11 years. Our highest

regard is 7 years. In the) period in question these birds have

been kept in the Western Aviary, with heated interior but access

to open air.

No. of Ay.dur. Max. dur.

BuceRoripa. Indiv. in months. in months.

Bucorvus abyssinicus (Ground Hornbill) 7 32 146

B. cafer (South African ,, ) 1 64. 64

Buceros lunatus (Lunated Hornbill) 1 Ott 17

B. rhinoceros (Rhinoceros 5 TSE 6 24. 75

Dichoceros bicornis (Concave-casqued 10 56 167

Hornbill)

Anthracoceros, whole genus ............... 9 24 51

A. coronatus (Crowned Hornbill)......... 4 25 48

A. malayanus (White-billed Hornbill). 1 10 10

A. malabaricus (Indian Pied 5 sian Al 26 51

Oranorrhinus corrugatus (Wrinkled- 1 1 1

billed Hornbill)

Aceros nepalensis (Nepalese _,, aa 5D 10

Oeratogymna elata (Klate a yd 48 138

Sphagolobus atratus (Black 53 5 40 12

Bycanistes subcylindricus (Subeylin- 2 59 94

drical Hornbill).

Rhytidoceros plicatus (Plicated Bi) pes 145 33

+ undulatus.

Toccus erythrorhynchus (Red-billed ,, ) 4 33 61

T. melanoleucus (Black and White ,, ) 2 47 58

The Hornbills are omnivorous, rather large birds, and the record

of maximum duration of 14 years for a Concave-casqued Hornbill

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 489

is by no means unexpectedly high. The relation of average

duration to maximum duration varies but is on the whole good,

showing a fairly high viability. These birds have been kept for

the most part in a large aviary with heated retreats but access to

the outer air.

No. of Av. dur. Max. dur.

Upupip®. Indiv. in months. in months.

Upupa epops (Hoopoe) ............0.2eseee 7 Wy 29

The maximum and average durations show low viability.

Momortip #.

Momotus brasiliensis (Brazilian Motmot) 1 74. 74

M. subrufescens (Cartagenian SNe tune” 98 121

The numbers of individuals are very small but would appear

to show a high viability for these omnivorous birds, which have

been kept in a heated aviary with open-air flight-cages.

MEROPIDA.

Merops apiaster (Common Bee-eater) ... 2 12 13

The very poor success with Bee-eaters is no doubt associated

with the difficulty of feeding them.

CoRACIIDE.

Coracias affinis (Allied Roller)............ 1 21 21

C.garrulus (Common ,, )........-... 6 6°5 10

The difficulty with these omnivorous but largely insectivorous

birds is probably the provision of suitable food.

PoDARGIDA.

Podargus cuvieri (Cuvier’s Podargus)... 4 12 32

There seems no reason except low viability to account for the

absence of success with this hardy-looking omnivorous bird.

STEATORNITHID.

Steatornis caripensis (Oil-bird) ......... 1 1 i

No just inference can be drawn from a single example.

No. of Ay. dur. Max. dur.

RHAMPHASTID&. Indiv. in months. in months.

Rhamphastos, whole genus............... 59 35 143

ies ariel: (Ariel Moucan) 2 3.20-...-+22 2 24 101

RR. brevicarinatus (Short-billed Toucan) 2 91 108

f. carinatus (Sulphur-breasted ,, ) 7 26 58

R. cuviert (Cuvier’s Se Maia 5 8

. dicolorus (Green-billed in ee 54 143

490 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Ay. dur. Max. dur.

RHAMPHASTID® (cont.). Indiv. in months. in months.

Rhamphastos erythrorhynchus (Red- 3 20 35

billed Toucan)

R. tocard (Doubtful Toucan) ............ 2 67 Jil

lieuoco (Moco Women) sssonsso20s00%2 9 44 93

R. vitellinus (Sulphur-and-white- 2 96°5 105

breasted Toucan)

Pteroglossus, whole genus ............+-. 20 29 87

P. inscriptus (Lettered Aracari) ...... 4) 20 30

P. torquatus (Banded ARS) PY Ys aie 6 30 tol

P. wiedi (Maximilian’s ,, ) ...... 9 32 87

Selenidera maculirostris (Spot-billed 2 20 25

Toucanet).

The Toucans and Aracaris are fruit-eating but partly omni-

vorous and flesh-eating birds, with simple intestines, the hind-gut

being relatively short and mot very capacious. Their viability as

shown by maximum and average durations is relatively good.

In the period in question they were kept in large cages in a

warmed house, without access to the open air.

Capironip a. Tue, fimoie, mans

Megalema, whole genus.................. 1 AO 122

M. asiatica (Blue-cheeked Barbet)... 6 A2 104

M. hodgsoni (Hodgson’s Tad) poe ee 2 4 4

M. virens (Great Die. ernie a 46 122

The Barbets have an almost omnivorous diet and have short

intestines, the hind-gut being relatively short and uncapacious.

Their viability as shown by maximum and average durations is

good, and in the period covered they were kept in cages in a

warmed house without access to the open air.

CucuLIp2. No.of Ay.dur. Max. dur.

Indiv. in months. in months.

Cuculus canorus (Cuckoo) ............+.. 21 7 38

Hudynamis orientalis (Black Cuckoo) 5 19 Al

H. taitensis (Long-tailed a ae) eal 3 3

Centropus, whole genus .................. 6 20°5 37

C. phasianus (Pheasant Coucal) ...... 1 19 19

C. rutipennis (Indian Serta enter 3 22 37

C. senegalensis (Senegal a yea 2 18°5 19

Oprationionigas (700 (C&GRI)) cceecceccscosenasece 9) 6 26

Guira piririgua (Guira Cuckoo) ...... 23 25 55

The Cuckoos are insectivorous and omnivorous birds with

relatively capacious hind-guts. Their viability as shown by the

maximum and average durations is low, but as is usual the failure

has been most complete in the case of the Common Cuckoo.

These birds have been kept under different conditions, sometimes

with, and sometimes without, access to open air.

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 491

No. of Ay. dur. Max. dur.

MusoPHacip®. Indiv. in months. im months.

Turacus, whole genus .............a.. 53 17°8 93

L. buffoni (Buffon’ Se NOUIACOUW)) se sans 5 13 20

7’. corythaix (White-crested Touracou). 9 28 93

7’. livingstoni (Livingstone’s se Se 11 14

T. macrorhynchus (Great-billed ,, ). 4 19 33

T. persus (Senegal ue er oe 15 60

T. schalowt (Bencuelay =). 37 37

Gallirex chlorochlamys (Green-necked 5 24 A5

Touracou).

G. porphyreolopha (Purple-crested 1 us 19

Touracou).

Musophaga violacea (Violaceous G 17 44

Plantain-eater).

The Touracous and Plantain-eaters are frugivorous and

omnivorous birds with relatively simple intestines, the hind-gut

being reduced. Their viability, as shown by maximum and

average durations, is not good but surpasses that of the Cuckoos.

They have been kept under varying conditions, always with

artificial heat, but in some cases with access to the open air.

Summary of Picarie.

Picarian birds form a heterogeneous group, united by no certain

or close bands of affinity, and it is not surprising that their

relative viabilities and longevities do not correspond with their

gradations of size in any close fashion, although there is evidence

of such a general relation, The maximum duration recorded is

nearly 14 years for a Hornbill; then come a Toucan with nearly

12 years,a Motmot and Barbet with 10 years,a Touracou with nearly

8 years, and a Kingfisher with 7 years. The conditions under

which these birds were kept during the period covered, have not

differed much. In the vast majority of cases they were kept in

houses or aviaries with artificial heat, but in some instances there

was access, at least occasionally, to open-air flight-cages. Iam

unable, however, to draw any clear inference with regard to the

effect of the mode of housing from the facts at my disposal. The

very marked differences in viability exhibited, I think, must

depend largely on real constitutional differences between the

different suborders. The diet of the whole set tends to be mixed,

with a strong leaning towards carnivorous or insectivorous

character. The character of the alimentary tract varies con-

siderably within the assemblage, and I thinka marked correlation

can be traced between the kind of gut and the viability. The

Caprimulgidz, Meropid, Coraciide, and Cuculide have all

relatively capacious hind-guts and ceca, and in all these the

average duration is low, generally under a year, whilst the maxi-

mum duration is never high. The Musophagide are generally

492 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

regarded as being closely allied to the Cuckoos; they have no

ceca and a relatively uncapacious hind-gut, and their average

duration is considerably over a year, whilst examples have lived

over 7 years. The Upupide have no ceca but a relatively

capacious hind-gut, and have a low viability. The Picide and

Alcedinide have no ceca; the hind-gut of the Woodpeckers is

certainly not specially capacious, and they must be regarded as

exceptions to the generalization; the Kingfishers are rather

better in viability, but in the family as a whole it is not good.

On the other hand, the Musophagide, as I have already men-

tioned, the Momotidx, the Coliide, the Capitonide, the Rham-

phastidee, and Bucerotide are all devoid of functional ceca, have

relatively short and uncapacious hind-guts, and show amongst

Picarian birds the highest average and maximum durations. The

group appears to furnish considerable evidence for Metchnikoff’s

generalization as to the relation between long duration of life

and the absence of opportunity for intestinal putrefaction.

It is interesting to note that Capt. Flower’s only high Picarian

record, that of a Coly still alive after 5 years’ duration, is in my

records surpassed by three different species of the same genus.

STRIGES.

STRIGIDE. Tabs | Sue ane

Stria flammea (Barn Owl)............... 67 17-4 60

Phodilus badius (Bay Wood Owl)...... 1 50 50

ASIONIDE.

HAO, WOO. SIMMS. Soosogsooscosassees96 000 ii 10 30

A. brachyotus (Short-eared Owl) ...... 22 11 30

A. mexicanus (Mexican Kared Owl)... 2 15 15

A. otus (Long-eared Owl). ............... 53 4 26

Syrnium, whole genus ...............--- 71 21 134

S.aluco (Tawny Ow)l)............... 60 18 AT

S. indranee(Indranee ,, )............... 4 55 134

S. nebuloswm (Barred ,, ).............4. 1 63 63

S. wuchale (Sharp's Wood-Ow))......... 2 20 28

S, sinense (Pagoda Owl) ......... 1 50 50

S. personata (Spectucled ,, ) ......... 1 12 12

S. uralense (Ural ya) Menten 1 2 3

S. woodfordi (Woodford’s ,, ) ......... 1 8 8

Bubvoey wholeteenus sence eee 88 32 yl

B. ascalaphus (Savigny’s Eagle-Owl)... 1 34 34

B. bengalensis (Bengal tS Pea 32 64

B. blakistoni (Blakiston’s ,, )... 2 16 23

B. coromandus (Coromandel ,, )... 1 29 29

L. lacteus (Milky a ee co 58 99

B. maculosus (Spotted aie We) eat italiy 27 59

B. magellanicus(Magellanic ,, )... 1 ls 7

B. maximus (Great pte) ae eS 28 81

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 493

No. of Ay. dur. Max. dur.

ASIONID& (cont. ). Indiv. in months. in months.

Bubo nepalensis (Nepal Eagle-Owl)... 1 108 108

B. orientalis (Orel me) Net 83 85

B. poensis (Fraser’s oF) Poems 45 107

B. virginianus (Virginian sin ane. LS 34°8 111

Ketupa ceylonensis (Ceylonese Fishing- 4 10 26

Ow 1).

K. javanensis (Javanese Fishing-Owl). 5 37 51

SCOMS a WNOLE GOMUS duane es cece ete seems 17 8°6 45

S. asio (American Scops Owl) ......... 1 26 26

egeU(SCOPS: Owill)k se. eeiss faeeca. anes 10 4:4 9

S. glabripes (Smooth-footed ScopsOwl) 1 45 45

S. lempiji (Horsfield’s & 1 1 1

S. leucotis (White-eared es Wakes 10 16

S. senegalensis (Senegal 3 ara 2 2

Scotopelia pelt (Pel’s Owl) ............... 1 5 5

Pulsatrix torquata (Downy Owl) ...... 10 40 87

Sceloglauax albifacies (White-faced Owl) 1 33 33

Athene brama (Spotted Owl) ............ 2 9°5 LZ

A, noctua (Naked-footed Owlet) ...... 14 13°5 34

Ninox boobook (Boobook Owl) ......... 3 26 50

NV. nove-seelandice (New Zealand Owl) 1 Ie iy

Speotyto cunicularia (Burrowing ,, ) 46 21 65

Glaucidium passerinum (Passerine ,, ) 13 6°5 17

G. phalenoides (Brown ,, 5 1 6 6

Nyctea scandiaca (Snowy Owl) ......... 9 22 75

Surnia funerea (American Hawk Owl) 1 5) 5

Summary of Striges.

¥Y g

We know from Mr. J. H. Gurney’s paper (doc. cit. p. 36) that

the potential longevity of Owls is far in excess of the maximum

durations recorded here. On excellent authority, that writer

cites an Eagle-Owl of 68 years, a Ketwpa of 39 years, and a

Tawny Owl of 26 years. The longest record on my list is that of

an Indranee Owl which lived for over 11 years in the Gardens,

and at the present time there are alive a Milky Eagle-Owl which

has been for 19 yearsand a Pel’s Owl which has been for 12 years

in the Gardens. Until comparatively recently the quarters

assigned to the Owls at the Gardens were extremely bad; it was

assumed that Owls required neither light nor space. They have

now been given better accommodation, and it is interesting to

state, that except the Barn Owls, all the Owls take the fullest

opportunity of sunning themselves and spend the greater part of

their time fully exposed. Although a good deal must be dis-

counted as a direct reaction to unfavourable conditions, it is still

clear that the viability of Owls is not good; not only are the

maximum durations far below what we know to be the potential

longevities, but the average durations are in most cases extremely

494 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

low. Owls, as is well known, have an unusually capacious hind-

gut with very large ceca.

It is interesting to notice that all Capt. Flower’s records of

Owls are relatively low, just over 3 years being the highest.

PSITTACI.

NESTORID2. Noe

Nestor notabilis (Mountain Ka-ka) ... 14

Lorip2.

Chalcopsittacus scintillatus(Red-fronted 1

Lory).

HOS, SANOE ANUS seageoasdsese-sooaseanosee" 13

EH. riciniata (Violet-necked Lory)...... 4

H. reticulata (Blue-streaked ,, )...... 3

JOST OMORED (ABE ICIS) \"aJocmaneonnnosesseson 4)

EL. wallacei (Wallace’s Lory) ............ 1

GCP OIE, WTO GENS Sarscsanossansseaeeo" 26

L. domicella (Purple-capped Lory)... 10

L. flavo-palliatus (Yellow-backed ,, _) 4

L. garrulus (Chattering ea etioat a

L. lory (Three-coloured ,, )... 7

L. tibialis (Blue-thighed ,, )... 1

Trichoglossus, whole genus............... 31

T'. cyanogrammus (Green-naped Lori- |

keet).

T. forstent (Forsten’s Lorikeet)...... 1

T. hematodes (Blue-faced ,, _ )...... 5

T.matehelli (Mitchel’s ,, )...-:: 1

T. nove-hollandice (Swainson’s Lorikeet) 14

T’. ornatus (Ornamental ))seee

T. rubritorques (Blue-bellied ,, ) 2

Psitteuteles chlorolepidotus (Scaly- 6

breasted Lorikeet).

Glossopsitiacus concinnus (Musky iat

Lorikeet).

CACATUIDA.

Microglossus aterrimus (Great Black 3

Cockatoo).

Calyptorhynchus banksi (Banksian 3

Cockatoo).

C. funereus (Funereal Cockatoo) ...... 2

Callocephalon galeatwm (Ganga Cockatoo) 1

Cacatua, whole genus................:..-: 84

C. alba (Greater White-crested 5

Cockatoo).

C. citrina (Citron-crested _,,

~——

C’. ducorpsi (Ducorps’s ‘

Av. dur.

in months.

nee

Gow nw =

Se Se or)

Max. dur.

im months.

227

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

Cacatums (cont.). ee

Cacatua galerita (Greater Sulphur- 8

crested Cockatoo).

C. goffint (Goftin’s Cockatoo). 5

C. gymnopis _—_ ( Bare-eyed Pe) hae

C. hematuropygia (Red-vented ,, ). 38

C. leadbeatert (Leadbeater’s ,, ). 17

C’. moluccensis (Rose-crested ,, ). 10

C. ophthalmica (Blue-eyed pre ace

G: roseicapilla (Roseate on) )aeelO

C. sanguinea (Blood-stained ,, ). ~~ 1

C. sulphurea (Lesser Sulphur- crested 8

Cockatoo).

C. triton (Triton Cockatoo) ............ 4

LInemetis nasica(Slender-billed Cockatoo) 5

LL. pastinator (Western ,, )

Calopsittacus nove-hollandice (Cockateel) 61

PsIrTaciDz&.

Anodorhynchus, whole genus............ 8

A. glaucus (Glaucous Macaw) ......... 1

A. hyacinthinus (Hyacinthine Macaw) 4

Am teams (dhean's: Wiacaw) s.a.0 95:2 e051. 2 3

Cyanopsittacus spixi (Spix’s Macaw) . 2

PAG Ww MNOle SeMUS es seg ehe eee eee 49

A. ambigua (Ambiguous Macaw)... 2

A. ararauna (Blue-and-Yellow ,, )... 16

A. chloroptera(Red-and- Yellow ,, _ ) 9

A. hahni (Hahn’s oe 2

A. macao (Red-and-Blue ,, )... 9

A, maracana (Illiger’s A eae ade

A. militaris (Military ah chee

A. nobilis (Noble eens) i

A. severa (Severe aks) 1

Conurus; whole genus... .56. denen: oo 90

C. acuticaudatus (Sharp-tailed Conure) 1

C. eruginosus (Brown-throated,, ) 24

C. aureus (Golden na

C. auricapillus (Golden-headed ,, ) 4

C. aztec (Aztec oa meee

C'. cactorwm (Cactus Ae de ane?

C. canicularis (Petz’s ao, ener

C. quarouba (Golden ue eee

OC. hemorrhous (Blue-crowned ,, ) 8

C. holochlorus (Mexican 2 had aan

C. jendayt (Yellow-headed ,, ) 12

C. nanday (Black-headed ,, ) 9

C. ocularis (Eyed Sel

C’. chloropterus (Green-winged ,, ) 5

Avy. dur.

in months.

66°8

113°5

AS*5

12:4

204

“I

Or Or

495

Max. dur.

in months.

113

182

131

496 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Av. dur. Max. dur.

PsITTACIDAL (cont.). Indiv. in months. in months.

Conurus rubritorquis (Red-collared 1 19 19

Conure)

C. rubrolarvatus (Red-masked ,, ) 1

C. solstitialis (Yellow a 4

C. wagleri (Wagler’s 5) eal 144 144

Conuropsis carolinensis(Carolina ,, ) 6

Cyanolyseus patagonus (Smaller Pata- 10

gonian Conure).

FHenicognathus enter viii (Slight- 1 184 184

billed Parrakeet).

Microsittace ferruginea (Chilian Conure) 1 36 36

Pyrrhura, whole genus .................. 21 54 172

P. cruentata (Red- eared Conure) ...... D 22 44

P. lewcotis (White-eared ,, )...... 9 51 112

P. perlata (Pearly A/F Meco d 2 98 172

P. vittata (Red-bellied spate Bt) ike 5 73 120

Myopsittacus monachus (Grey-breasted 15 40 124

Parrakeet). .

Bolborhynchus lineolatus (Lineolated 4 34 48

Parakeet).

Psittacula passerina (Passerine Parrot) 17 22 57

Brotogerys, whole genus.................. 30 3D 93

B. chrysopterus (Golden-winged 1 57 57

aie

B. jugularis (Tovi 5 34 65

B. pyrrhopterus( Orange- flanked , pal a cea 52 78

B. tirica (All-green Le NE GOS 32 93

B. twi (Tui Bi ane) 30 46

B. tuipara (Golden-fronted ,, ) 1 45 45

B. virescens (Yellow-winged A yo 2 19 20

Chrysotis, whole genus) =)5...-2-----.-.): 168 43 312

C. estiva (Blue-fronted Amazon) 11 36 103

C. agilis (Active FoR 3 i

C. albifrons (White-browed ,, ) 10 18 74

C. amazonica (Orange-winged ,, ) 9 55 228

C. augusta (August SIN Rad oe 359 70

C. auripalliata (Golden-naped ,, ) 8 83 312

C. autumnalis (Yellow-cheeked ,, ) 8 34 122

C. bodint (Bodinus’ ioe) irae 83 153

C. bouqueti (Bouquet’s seth iacuae 72 72

C. brasiliensis (Red-tailed Ake) Veet 66 128

C. dufresniana (Dufresne’s BY Ne) aes 20 35

C. caymanensis (Cayman ia) i it

C. collaria (Red-throated Be TAO 36 62

C. farinosa (Mealy Be He AG 52 68

C. festiva (Festive Be NO aa 35 AQ

C. finschi (Finsch’s ee) aes 63 121

C. guatemale (Guatemalan Bee Yoga 14 28

C. guildingi (Guilding’s oe) Al 69

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

No. of Ay. dur.

PSItraciD® (cont.). Indiv. in months.

Chrysotis leucocephala (White-fronted 8 32

*Amazon)

C. levaillanti (Levaillant’s 5 ee Ss 68

C. mercenaria (Mercenary 5 aie 13

C. ochrocephala (Yellow-fronted ,, ) 8 45

C. ochroptera(Yellow-shouldered ,, ) 6 50

C. panamanensis (Yellow-billed ,, ) 6 60

C. pretrit (Prétre’s epee 12

C. rhodocorytha (Red-topped ow TRG 36°5

C. salwina (Salvin’s Bey) eG 40

C. ventralis (Sallé’s ot ye ee 42

C. versicolor (Blue-faced spi 25

CO. vinacea (Vinaceous FN) 34

C. viridigena (Green-cheeked Witaae tele) 63°5

C. vittata (Red-fronted Pe a ae 48

C’. cantholora (Yellow-lored thea’ MRS 46

Pachyurus brachyurus (Short-tailed 2 30°5

Parrot).

TEUOOUIR NANO (HNCEY Son goagcoaqsseapennes 28 29

Vig chalcopterus (Bronze- -winged Parrot) 2 26

P. fuscus (Dusky ta) 9 14°5

P. maximiliant (Maximilian’s ,, ) 3 69

P. menstruus (Red-vented cl 23°8

P. senilis (White-headed ,, ) 1 id

P. sordidus (Sordid }) 59)

Deroptyus accipitrinus (Hawk- headed 9 49

Caique).

Pionopsittacus pileatus (Red-capped 2 1

Parrot).

Carvcan whole! genus 2.24. 2:00. yh eee 16 AT

C’. leucogastra (White-bellied Caique). 2 6°5

C. melanocephala (Black-headed ,, ). 12 39

C. xanthomera (Yellow-thighed ,, ). 2 130

Peocephalus, whole genus ............... 46 32

P. fuscicapillus(Brown-headed Parrot) 2 53°

P. fuscicollis (Brown-necked pa as a4) 72)

P. gulielmi (Jardine’s 52) 12

P. meyert (Meyer's ie) yy aS AD

P. robusius (Levaillant’s yet ro be 2 24

P. rueppelli (Riippell’s ene Im HEG 3°8

P. senegalus (Senegal hdd ingle) 30

Psitiacus erithacus (Grey Ea NEDO, 13

P. timneh (Timneh ASA AL 84

Coracopsis, whole genus................5- 13 164:7

C. barklyi (Praslin Parrot)............ 6 93

Criigra) (lesser Vasae 5... ))\eeesese ea: 4 98

C. vasa (Greater ,, a5, eM hots, 3 397

Ficlectus, whole genus..................0+: 35 31

E. cardinalis (Cardinal Kelectus) ...... 1 38

497

Max. dur.

in months.

498 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Avy. dur. Max. dur.

PsITTactDz (cont.). Indiv. in months. in months.

Eclectus pectoralis (Red-sided Kclectus) 13 34 90

E. roratus (Grand pees) elt AT 119

E. westermani (Westerman’s ce DRS A-6 15

Tanygnathus, whole genus ............ 12 62 189

T. luzonensis (Blue-crowned Parrakeet) 3 19 37

T. megalorhynchus (Great-billed ,, ) 1 154 154

T. muellert (Mueller’s Sl) aS 66 189

Paleornis, whole genus ...5.0-4,.---4-.- 96 4] 250

P. alexandri (Alexandrian Parrakeet). 15 53 247

P. caniceps (Grey-headed - Spy 2 2

P. cyanocephala (Blossom-headed ,, ). 11 56 DT

P. docilis (Rose-ringed _ ,, ) 8 A3 157

P. fasciata (Banded b5 ye ue 69-6 160

P. longicauda (Malaccan % ) itanael 12 26

P. nicobarica (Red-cheeked ,, ) 1 3 3

P. peristerodes (Malabar i )fioerl AT 86

P. rosa (Rosy ai ) 6 15 30

P. torquatus (Ring-necked _,, ee) 34 250

Patuygeles. witOle eeu sites pecan eee eneee 15 31 72

P. alexandre (Princess of Wales’ 3 23 AQ

Parrakeet)

P. barrabaudi (Barrabaud’s ss ) 4 yes 12

P. melanurus (Black-tailed - es 23 AO

Ptistes erythropterus (Red-winged 13 33 61

Parrakeet).

Aprosmictus coccinopterus (Smaller 1 162 162

Red-winged Parrakeet),

A. cyanopygius (King 3 ar 1) 27:5 80

Pyrrhulopsis, whole genus............... 17 A4 116

P. personata (Masked Parrakeet) ...... 7 52 89

P. splendens (Shining 5 th be 5 44 116

P. tabuensis (Tabuan i Meaect 5 3 133

Psittinus imcertus (Blue-rumped 1 20 20

Parrakeet).

Agapornis, whole genus .................. 68 23 oi

A.cana (Grey-headed Love-bird)... 17 30 97

A. pullaria (West-African ,, )... 38 16 56

A. roseicollis (Rosy-faced sie pie te 22 40

LEorculus, whole genus) se-)..40ess seer 40 Ie 80

L. chrysonotus (Golden-backed 3 26 31

Hanging-Parrakeet).

L. galgulus (Blue-crowned Hanging- 21 ly 80

Parrakeet).

L. indicus (Ceylonese Hanging- 15 15 Bee

Parrakeet).

L. pusillus (Javan Hanging-Parrakeet) 1 9 9

Platycercus, whole genus ............... 83 42°6 202

P. adelaide (Adelaide Broad-tail)...... 2 31 52

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

PsrrraciD& (cont.).

Platycercus barnardi(Barnard’s Broad-

tail)

P. browni (Brown’s Broad-tail) ...

P. elegans (Pennant’s ,, ae

P. erythropeplus (Red-mantled "Broad

tail).

P. eximius (Rose-hill Broad-tail) ......

P. flaveolus(Yellow-rumped Broad-tail)

FP, flaviventris (Yellow-bellied ,, ,, )

P. icterotis (Stanley Broad-tail).........

P. pallidiceps (Pale-headed Broad-tail)

P. pileatus (Pileated ih oe)

P. semitorquatus (Yellow- collaréd

Broad-tail).

P. zonarius (Bower's et ee)

Psephotus, whole genus .....:............

P, chrysopterygius (Golden-winged

Parrakeet).

P. hematonotus (Blood-rumped

Parrakeet).

P. multicolor (Many-coloured ,, _ ).

P. pulcherrimus (Beautitul et)

Weophema, whole genus'..........-.+:-...

NV. bourkii (Bourke’s Grass-Parrakeet)

LV. chrysogastra (Orange-bellied Grass-

Parrakeet).

NV. elegans (Klegant Grass-Parrakeet)

N. pulcheia (Lurquoisine Grass-

Parrakeet).

NV. splendida (Splendid Grass-

Parrakeet)

N. venusta (Blue-banded Grass-

Parrakeet).

Cyanorhamphus, whole genus .........

C’. auriceps (Golden-headed Parrakeet)

C. matherbi (Alpe

C. nove-zealundie (New Tiealana

Parrakeet),.

C. saisseti (Saisset’s oe ae

C’. unicolor (Antipodes Island ,, )

Nymphicus cornutus (Horned ,, ).

NV. weeensis (Uvean ,, :

Nanodes discolor (Swift bs

Melopsittacus wndulatus (Undulated

Grass-Parrakeet).

Geopsittacus occidentalis (Western

Ground-Parrakeet),

STRINGOPIDA.

Stringops habroptilus (Night Parrot) .

No. of

Indiv.

bo BR co bp & bo

OO je

Ewe

Ay. dur.

in months.

A99

Max. dur.

in months.

202

500 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

Summary of Psittaci.

Popular repute and exact records agree in assigning a high

potential longevity to Parrots. Metchnikoff (op. cit. p. 52) cites

on the authority of Levaillant a Grey Parrot of 93 years, on that

of Butler, a Sulphur-crested Cockatoo of 81 years, on that of

Abrahams, an Amazon of 102 years, whilst he himself owned an

Amazon which died at an age between 70 to 75 years. ‘The

figures in Mr. J. H. Gurney’s list (op. cit. p. 36) are lower, the

highest being a Blue Macaw of 64 years. ‘The figures of

maximum duration at the Gardens are very much lower in most

cases. A Greater Vasa Parrot (already cited by Gurney) lived

for 54 years. With regard to the others the figures of maximum

duration are distributed nearly equally throughout the different

groups of Psittaci; thus an Amazon and a Banksian Cockatoo

reached about 26 years, a Macaw 23 years, a Conure, a Poeocepha-

lus Parrot anda small Parrakeet 20 years, a Vestor 19 years,a Lory

18 years, and a Lorikeet 17 years, whilst there are at present

alive a Lesser Vasa Parrot which has lived in the Gardens for

26 years, and a Lorikeet of 17 years’ duration.

I have no information as to the ages of the Parrots on their

arrival at the Gardens, but as in an assemblage of 1480 birds,

selected at random so far as age is concerned, there must have

been many young birds, the maximum ages attained would

appear to show that Parrots have not a high viability under the

conditions to which they have been subjected, as otherwise there

would have been a more frequent and closer approach of maximum

duration to potential longevity. The records of average duration

are not more favourable. The great majority of the birds had an

average duration of somewhere between 2 and 4 years, a few such

as the Lorikeets, Cockatoos, Conures, and Caiques were in most

cases between 4 and 6 years, whilst the Macaws usually showed

averages of over 6 years. I can see only the vaguest possible

correlation between average and maximum durations and size, but

on the whole there are more of the larger birds amongst the

higher figures.

As we know that Psittaci have a high potential longevity, it

is clear that either the conditions under which they are kept in

captivity are markedly unfavourable, or that these birds have a

low viability. Im the period in question, the examples of this

group were kept in a rather crowded house, well warmed in cold

weather. A certain number of them were placed out of doors

(chiefly Macaws and Cockatoos) in good weather, but were

chained to perches. The vast majority of them were in relatively

small cages, and never had access to open air, whilst none had

the opportunity of flying. It is of course clear that exceptional

individuals can do well without exercise, and it is highly

probable that at least most of the birds cited as instances of high

longevity, were kept in cages or on stands. But it is equally

clear that such conditions, and in particular the absence or fresh

air, are not favourable.

RELATIVE VIABILITY IN MAMMALS AND BIRDS. DOL

Parrots are typically vegetarian birds, although, as is well

known, most of them do well if some animal material is added to

their diet. The intestinal tract is long and unusually complex, but

it is narrow in calibre, there are no ceca, and the greater portion

appears to correspond with the small intestines, whilst there is

little that is used as a reservoir for waste material, or that can be

a probable seat of putrefaction. I cannot see that the indications

point towards their low viability being the result of their consti-

tution, and am disposed to attribute the relatively low maximum

and average durations to unsatisfactory conditions. The Zoolo-

gical Society has made experiments in recent years, in giving the

larger parrots free exposure to the open air, without artificial

heat. The experiment has not been of sufficiently long duration

to test the effect on longevity; but even though the conditions

are not specially favourable in the particular aviary, the general

result appears to be good. JI have no doubt that the ideal

management is a combination of open-air flight cages with

sheltered, and quite possibly warmed retreats.

Capt. Flower’s: best records are a Bare-eyed and a Slender-

billed Cockatoo, still alive after 12 years’ duration. As it was

only in 1898 that he took char ge of the Giza Gardens and began

to make his records, about 12 years’ duration was the maximum

possible.

ACCIPITRES.

peor Dy, inion, in mouthe

Pandion haliaétus (Osprey) .,..,....... 15 16 84.

Open smnOleGeNUS. a... +s. -nrdecuan: =. « 23 15 43

C. eruginosus (Marsh Harrier) ...... 5 1) 42

C. cineraceus (Montagu’s ,, ) ...... 9 9 23

C. cyaneus (Hen A Dh anheeee Baal 10 10

C. gouldi (Gould’s Sn) apes 1 14 14

Comacrunus (Swamson’s, ,,) ..... _ L A3 4R

C. maurus (Moor A eo ibeus ic 6 16 oe)

BULCOn NIMOIS) SEMUS. ) sxeeg nsseoce se cates 95 25 225

B. albicaudatus (W hite- tailed Buzzar d) 4 AZ 116

B. auguralis (Augura RM 8:5 13

B. borealis (Red-tailed an ea lel 24 93

B. desertorum (African a BIO) 37°6 pia

B. erythronotus(Red-backed ATES) wines 18-5 3

B. ferox (Long-legged eis) 16 20

BL. jackal (Jackal Satie?) es 34 81

B. pennsylvanicus(Pennsylvanian,, ) 2 30°D Al

&. poliosomus (Patagonian ee) en 25 25

B. vulgaris (Common ) 42 19-5 72

Archibuteo lagopus (Rough- lege ged 22 14:5 36

Buzzard).

Pernis apivorus (Honey Buzzard) ... 4 57 122

P. ptilorhyncha (Crested Honey Buzzard) 1 22 22

Proc, Zoou. Soc.—1911, No. XX XIII, 33

502 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

FALconrD& (cont.).

Haliastur indus (Brahminy Kite)

H. intermedius (Javan Brahminy Kite)

Asturina, whole genus ............+..0+-

A. magnirostris (Great-billed Buzzard

Hawk).

A. nattereri (Natterer’s Hawk) .........

A, nitida (Shining Buzzard Hawk) ...

Urubitinga, whole genus ............--.

U. anthracina (Ash-coloured Uru-

bitinga).

U. meridiondlis (Rusty Urubitinga)...

U. zonura (Banded-tailed Urubitinga)

Buteogallus nigricollis (Black-necked

Buzzard).

Leucopternis palliatus (Mantled

Buzzard),

Geranouétus melanoleucus (Chilian

Sea-Hagle).

Harpyhaliaétus coronatus (Crowned

Harpy).

Morphnus guianensis (Guianan Crested

Eagle).

Thrasattus harpyia (Harpy KHagle)...

Helotarsus ecaudatus (Bateleur ,, ) ...

Haliaétus, whole genus .........5......5.

H. albicillus (White-tailed Eagle)......

H. leucocephalus (White-headed Sea-

Eagle).

H. leucogaster (White-bellied Sea-

Eagle).

H. leucoryphus (Mace’s Sea-Hagle) ...

H. vocifer (Vociferous ,,_ ,, pel

Polioaétus plumbeus (Plumbeous Fish-

Kagle.)

Aquila, whole genus ............2-.-...--

. adalberti (Prince Adalbert’s Kagle) .

. audax (Wedge-tailed :,

. chrysaétus (Golden mit)

.clanga (Spotted e

. imperialis (Imperial te Palys

. nevioides (Tawny )

. verreauxt (Vulturine )

Nisaétus, whole genus ..................

NV. fasciatus (Bonelli’s Eagle) .........

N. pennatus (Booted 5s: At) edtco aecpen

WV. spilogaster (African Hawk-Kagle)...

Spizaétus, whole genus ..................

S. bellicosus (Martiai Hawk-Hagle) ...

bf Re he fa he

No. of

Indiv.

)

pe OD

Ay. dur.

in months.

62°5

116

2°5

Max. dur.

in months.

110

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

Fatconip# (cont.). Hes oF

Spizaétus caligatus (Malayan i

Hawk- Eagle).

S. ceylonensis (Ceylonese ,, ,, )... 3

S. coronatus (Crowned RPP Hg i) 10

S. wipalensis (Spotted Ast ee ee

S. orientalis (Japanese ,, ,, )... 1

S. ornatus (Ornamented,, ,, ) 2

S. tyrannus (Tyrant Me), on) ane 1

Lophaétus occipitalis (Black-crested 5

Kagle).

Circaétus gallicus (Short-toed Eagle) . 13

Dryotriorchis spectabilis (Beautiful 1

Wood-Hawk).

Spilornis, whole genus ..............06++ 4

S. bacha (Bacha Eagle) ................+. 2

mencela (Cheela,) 43.) sie edsincceewntete 1

S. spilogaster(Ceylonese Serpent-eater) 1

Acewpiter, whole genus ................+. 11

A.nisus (Sparrow Hawk) ............ 9

A. pileatus (Pileated ,, ) ............ il

A. melanoleucus (Black and White 1

Hawk).

Melierax, whole genus ............0..... 12

M. = monogrammicus (One-streaked 7

Hawk).

M. musicus (Chanting Hawk)......... il

M. polyzonus (Many-zoned ,, )......... 4A

Astin. Whole SCNUS .2.3.22.52.-cscles tact 35

A. approximans (Allied Goshawk)... 4

A. nove-hollandic (White a a) orale

A. palumbarius (Common ,, )... 22

A. tachiro @lachiromee ew) see

A, tibialis (Gambian yam )e. > ol

Paleo, wholevyeenus) ..5 5... sees vests coe 108

EN OSaLOTN INVER eee ton Woohoo ce ete 7

fF, biarmicus (Bearded Falcon) ...... 1

Ff. concolor (Ash-coloured ,, ) ...... 2

fF. eleonorce (Kleonora ata ee SAFE or 1

LF, feldeggi (¥eldege’s at Nemec 1

Ff. fusco-cerulescens (Orange-thighed 2

Falcon).

F, lanarius (Lanner Falcon) ............ 14

FF. melanogenys (Black-cheeked Falcon) 5

F. peregrinus (Peregrine Falcon) ...... 61

F, punicus (Mediterranean Peregrine 1

Falcon).

imsacer (Saker Walco), .2.:-cs-acascee- 3

iesubbuieo (Hobby) | <.02-dsgesedeoy ss 12

Hierofalco candicans (Gr eenland Falcon) 2

Ay. dut.

in months.

—O Ore

me wenden

oO Kop

—

He ete OUND GO OO

Or Or

503

Max. dur.

in months.

fiw

He RR oN ON

boN Ol Or bo 6

KE ONMWOIoc Ore =

504 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

FALCONIDE (cont.). Ne. oF

Hierofalco islandus (Iceland Falcon)... 6

Hieracidea berigora (Berigora Hawk). 5

Tinnunculus, whole genus ............... 81

T. alaudarius (Common Kestrel)...... AD

T’.. cenchris (Lesser in) eat 9

T. cenchroides (Nankeen ,, )...... 2

T. dominicensis (Dominican ,, )...... 1h}

T’. gracilis (Slender se ue) yacans 2

T. moluccensis (Molucca ,, _)...... 1

1 (royadoolins (Sky Ibe 4, onnee 2

T. sparverius (American .,, )...... 4

T. vespertinus (Red-footed ,, )...... 3

Hypotriorchis chicquera (Red-headed 1

Merlin).

Milints wi holey ce muse neeece eee met Ger 30

M. govinda (Indian Kite)............ 4

M. migrans (Black ME) ee err 8

M. egyptius (Egyptian ,, )............ 12

iM michinus, (Common ii) ))ee eer 6

Baza lophotes (Crested Black Kite) .. I

Hlanus ceerwleus (Black-shouldered 6

Kite).

Rostrhamus sociabilis (Sociable Marsh- 2

Hawk).

Herpetotheres cachinnans (Laughing 2

Falcon).

Milwago, whole genus ...................-. 26

M, australis (Forster’s Milvago) .. 5

M. chimachima (Chimachima ,, )... 5

M. chimango (Brown Lorie aL

M. megalopterus (Long-winged ,, )... 2

Polyborus brasiliensis (Brazilian 7

Caracara).

Polyboroidestypicus(Banded Gymnogere) 2

VULTURIDE.

Valiiirmewholereenus a reeestrenereekeer eter ite

V. auricularis (Sociable Vulture) ...... 7

V. calvus (Rondichernyar eee 3

V. monachus (Cimereous ,, _) ...... 8

V. oceipitalis (Occipital ,, ) ...... 1

Gos awihOlenOOMus teers. scan. ee see ieeee 14

G. bengalensis (Bengal Vulture) ...... 3

G. fulvus (Griffon WAP) 5 ASR 8

G. kolbi (Kolbe’s’ BSL Oe teee 1

G. rueppelli (Ruppell’s ,, ) ...... 2

Neophron percnopterus (Kgyptian 9

Vulture).

Ay. dur.

in months.

Max. dur.

in months.

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 50d

VULTURIDE (cont.). No.of — Av. dur. | Max. dur.

Indiv. in months. im mouths.

Gypohierax angolensis (Angolan 12 40 66

Vulture).

Gypattus barbatus (Kuropean Bearded 6 118 284

Vulture).

SERPENTARIIDE.

Serpentarius reptilivorus (Secretary 26 12 59

Vulture).

CATHARTIDS.

Catharies, whole genus ............6...5. 23 22 70

C. atratus (Black Vulture) ............... 15 18 57

Caaura (Turkey, os) msgs ence ee fi 30 70

C. urubitinga (Yellow-headed Vulture). — 1 8 8

Sarcorhamphus cequatorialis (Brown 1 96 96

Condor).

Seagua pies: (Condon)i icra nase. oer = 6 155 403

Gypagus papa (King Vulture) ..!...... 18 20 87

Summary of Accipitres.

The potential longevity of Accipitrine birds is well known to be

great, although, as in most other cases, well-authenticated records

are considerably lower than the ages assigned by popular rumour.

Metchnikoff (op. cit. p. 53) refers to a White-headed or Egyptian

Vulture of 118 years, and a Golden Hagle of 104. Mr. J. H.

Gurney’s exact records (loc. cit. p. 37) range downwards from a

Bateleur Eagle of 55 years and a Condor of 52. The maximum

durations in my record are considerably lower; those over 20 years

are a Caracara of over 41, a Condor of nearly 34, a Chilian Sea-

Kagle of nearly 28, a Sociable and an Angolan Vulture which

reached nearly 24, a Griffon of nearly 22 and a Wedge-tailed

Eagle of more than 21 years; whilst there is at present alive, a

Milvago that has been over 24 years in the Gardens. The

average durations mostly under 2 years are of Ospreys, Harriers,

Hawks, Falcons, Kestrels, Secretary Bird, and the American

Vultures; those mostly between 2 and 4 years, are of Buzzards,

Sea-Hagles, and the larger Hawk-EHagles; those mostly between

4 and 6 years are of the Chilian Sea-Hagles, Goshawks, and Kites ;

whilst the records above 6 years belong chiefly to Wedge-tailed

Eagles, Imperial Eagles, the larger Vultures and the Condor.

There is a rough relation between size and viability, and there is

the usual very low viability of the common European forms.

The conditions under which the birds of prey have been kept,

in the period covered, cannot be said to have led to a favourable

result. In the great majority of cases no artificial heat was

provided, even in winter, and there was full access to open air,

but especially in the case of the smaller birds, the quarters were

small and dark, and there was no provision for exercise. If the

records be compared with those of the Striges, it will be seen,

506 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

however, that both average and maximum durations are con-

siderably better in the case of the diurnal birds of prey. No

intelligent anatomist can now doubt that the two groups are far

separate amongst birds, and one of the notable differences is in

the structure of the alimentary tract. In all the Accipitrme

birds, although the intestines are relatively long and arranged in

a complex pattern, the hind-gut and cca are extremely reduced,

and the portion of the gut in which there is the greatest

opportunity for intestinal putrefaction therefore has a relatively

very small capacity.

Capt. Flower records only two Accipitrine birds—a Griffon

Vulture and an Egyptian Vulture—that are still alive after

12 years’ duration, the maximum possible in his records.

STEGANOPODES.

POLL 8 eM ahs ost

Pelecanus, whole genus .................. 21 139 380

P. consprciliatus (Australian Pelican). 2 285 380

P. crispus (Crested st a) de icra 274°5 329

P. fuscus (Brown Viet as 65 121

P. manillensis (Spot-billed ,, ). 2 54 65

P. mitratus (Mitred 5 ss) an 8) 130 261

P. onocrotalus (White Shea actin 239 300

P.rufescens (Red-backed ,, ). 6 66 152

Mr. J. H. Gurney (loc. cit. p. 38) gives the records. of six Pelicans

ranging from 41 to 18 years, and the maximum durations given

above, ranging downwards from nearly 32 years, are all re-

markably high, whilst the average durations show that these

birds are probably the most highly viable of all the birds and

mammals usually kept in captivity. They were not provided

with artificial heat, but throughout the day were exposed fully

to the weather, whilst at night, and occasionally in very cold or

foggy conditions, they were shut up in « small wooden shelter.

As compared with other fish-eating birds, the alimentary tract 1s

relatively short and simple, and the ceca and hind-gut are small

(although not so reduced as in Accipitrine birds) and uncapacious.

Capt. Flower records one White Pelican still alive after

12 years.

No. of Ay. dur. Max. dur.

PREGATIDE. Indiv. in months. in months.

fregata aquila (Great Frigate-bird) ... 5 45:6 91

It is most probable that only weak or injured examples of this

oceanic bird usually find their way into captivity, and the records,

t think, must be interpreted as showing a high viability. The

intestinal tract is simpler than that of Pelicans, with the hind-

gut and ceca more reduced. These birds have been kept in an

aviary with warmed interior and open-air front cages.

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 507

No. of Ay. dur. Max. dur.

PHALACROCORACIDS. Indiv. in months. in months.

Sula, whole. Genus: |... b.sciechd estar arenes 39 8°3 AQ

Se Omssaqeie (CANN) cere ctsaoets nehines he oh 27 6 19

S. leucogastra (Brown Gannet) ......... 5 15 40

S. piscator (Wines acre Nees a 2 14 19

S. serrator (Australian ,,- ) ......... 5 8°8 16

Phatacrocorax, whole genus.....:......... 84 12 94

P. brasilianus (Brazilian Cormorant)... 9 10°5 5D

P. africanus (African 35 ) ee 23° 36

P. carbo (Common - }) Bae e331 10°5 52

P. carunculatus (Rough-faced ,, ) 2 28 49

Pe AGROCULUS (SWAL) 556 sisnctaecSvenese te 15 10 44

P. javanicus (Javan Cormorant) ......... 1 3 38

P. lugubris (Mournful __,, Dear 1 94 94

P, varius (Pied i yh iakat 1 1 ih

Mr. J. H. Gurney (loc. cié. p. 38) quotes from H. O. Forbes

the record of a Common Cormorant which lived for 23 years.

The maximum durations on my list, ranging from nearly 8 years

downwards, are very much lower, whilst the average durations

of both Gannets and Cormorants seldom exceed one year. There

is the usual remarkable failure with local species—a failure which

would appear much greater if a considerable number of entries of

birds that lived less than a month had not been excluded from

the records. The intestinal tracts are of the fish-eating type—

relatively large and of smail calibre, but the hind-gut and ceca

are very rudimentary, and there seems little opportunity for

intestinal putrefaction.

No. of Av. dur. Max. dur,

PLOTID. Indiv. in months. in months.

PialohuseWwiOle SeNUS| Jreseut eee! see. 16 22 103

Plotus anhinga (American Darter)...... 12 18 53

P. levaillanti (Levaillant’s ,,_)...... 2 18 34.

P. melanogaster (Indian oy) 2 55°5 103

J. H. Gurney (loc. cit. p. 38) gives an instance of an American

Darter of 12 years old; I do not know of any other records.

The intestinal tract is similar to that of other Steganopodes, but

although the caeca (or single cecum) are vestigial, the hind-gut

is relatively more capacious and larger. These birds were kept

in a house (the Fish House) that was warmed in winter, and

their average duration and maximum duration show no high

viability.

Summary of Steganopodes.

Pelecans, Frigate-birds, Gannets, Cormorants and Darters,

whether or no they form a coherent zoological group, differ very

little in their requirements in captivity, and receive very much

the same treatment. ‘They appear to me to be a group of potentially

508 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

great longevity and viability, these, moreover, being fairly in pro-

portion tosize. Therecords of Phalacrocoracide are much lowered

by the number of European birds, which like most European

animals have a very heavy death-rate in captivity.

HERODIONES.

: v. dur. Max. dur.

a Iain, Rete in months.

Ardeu, whole emus’ Aee-nes esse eens 90 25 110

A. agame (Agami Heron) il 4) 5

A,alba (Great White ae) 4 TOD 26

A. atricollis (Blaick-necked 5 7) 2. 24 24

A. bubulcus (Buff-backed ,, )... 4 50 64

A. candidissima (Snowy sae) 10 20 74.

A. cinerea (Common ne) 23 28 SS

A. cocor (Cocoi pe 4 A2 93

A. egretta (Great American Eer et). 16 ol 100

A. garzetta (Little AUAK: ania 3D 98

A. goliath (Goliath Hieron)............... 1 30 35

A. gularis (Slaty ble) aceite tyes 7 19 72

A, herodias (Great American Heron). 1 31 31

A. nove-hollandice (White-faced ,, ). 2 13 Ne

A. purpurea (Purple aaah) Satna) 4-6 10

A. ralloides (Squacco Hane ee 6:5 Il

A. sumatrana (Typhon pe Bea 69 69

Butorides, whole genus .................. a 14 53

B. atricapillus (Blue-capped Bittern) . 2 44-5 D3

L. cyanurus (South American Little 1 12 12

Bittern).

B, virescens (Green Bittern) ............ 6 4 13

Ardetta involucris( Variegated Bittern). 3 23 40

A. minute (Mittle Bittern)............... 8 6 19

Nycticorax, whole genus.................. 64 30 199

NV. caledonicus (Nankeen Night 2 99 107

Heron).

NV. gardeni (Garden’s Night Heron). 6 9 32

N. griseus (Common _,, ate) te val 37 199

iV. violaceus (Violaceous 3 eh AVS TLS 19°5 61

Cancroma cochlearia (Boat-bill)......... ~) 21 53

Botaurus lentiginosus (American 1 8 8

Bittern).

JB Al Gres (ASNT) 40.5 y2houskoasesco5es 3 23 54

Tigriosoma brasiliense (Tiger Bittern). 6 48:9 a wullo2

1’. leucolophum (White-crested ,, ,, ). 7 20°5 33

J. H. Gurney (loc. cit. p. 38) gives somewhat doubtful records

of 60 and 30 years, and a well authenticated record of 22 years for

a Common Heron. <A duration of over 16 years for a Night

Heron is the best on my list. The average and maximum

Pox

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 509

durations show a rough correspondence with size; but the

relations of average to maximum duration show rather a low

viability for these birds. The intestinal tract is of the fish-eating

type, relatively very large and of small calibre, but with vestigial

ceca and with a reduced hind-gut. The conditions under which

these birds have been kept differ; some were placed in open

enclosures, others, especially in winter, in an aviary with warmed

interior.

CICcONIIDA. No. of Ay. dur. Max. dur.

Indiv. in months. im months.

Ciconia, whole genus ...........0.6.0:: 56 25 111

Csaba (OV biteStorls)\2.. 2.02... 34 26 74

Olboyciana(Boyce’s: ,, )......5.0--.-4- 2 14-5 28

Ceuiord = (Black, —0)) sees eee 20 20 ital

Dissura episcopus (White-necked 12 30 64

Stork).

D. maguari (Maguari Stork) ......... 10 40 84

Abdimia sphenorhyncha (White- 3 D0 81

bellied Stork).

NXenorhynchus australis (Black-necked 11 18 68

Stork).

X. senegalensis (Saddle-billed ,, ). 4 26 59

Mycteria americana (American 8 34°59 6d

Jabiru).

Leptoptilus, whole genus ............... 19 46°5 227

L. argala (Indian Adjutant) ......... 4) 17 227

L. crumeniferus (Marabou Stork) ... 9 47 NLS

L. javanicus (Javan Adjutant) ...... 5 15 35

Scopus umbretia (Tufted Umbre) Dd 24 43

Tantalus loculator (American 7 105 23

Tantalus).

Pseudotantalus ibis (African ,, ). 13 15 onl

P. leucocephalus (indiang) 9,0): jets & ness 23

Mr. J. H. Gurney (loc. cif. p. 38) cites well authenticated

records of a Black Stork of 30 years and a Jabiru of 36 years.

Nearly 19 years for an Adjutant is the highest on this list,

whilst the average durations vary, roughly in proportion to size,

from about 4 years to about | year. During part of the period

in consideration, the larger tropical Storks were kept m the

Antelope House, which was warmed in winter; towards the end

of the period a new house was erected, in which there were

heated compartments leading to open-air paddocks to which the

birds were admitted at the discretion of the keepers—a discretion

usually exercised with a strong bias in favour of heat as opposed

to air. The intestinal tract is relatively long and of small calibre

but the hind-gut and ceca are very reduced. There appears to

be no good reason in the structure of these birds for the rela-

tively low viability which appears in the records.

510 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Avy. dur. Max. dur.

PLATALEIDE. Indiv. in months. in months.

Platalea alba (African Spoonbill) ...... 2 21 29

P. leucorodia (Spoonbill) ............... ky) 16°5 115

Ajaja rosea (Roseate Spoonbill)......... ial 19 38

Tots ewnoleys ens seen eee eee carers if 80 220

I. ethiopicus (Sacred DIG) cere teense. 8 94 1G

i Germieng (SCRMIeti Sh eae Pere reneerere 2 25 38

I. melanocephalus (Black- -headed Ibis). 1 83 83

I. strictipennis (Australian Sacred ,,). 6 87 220

Carphibis spinicollis (Straw-necked 4 109°8 300

Ibis).

Nipponia temmincki (Nippon ,, ). 1 5 4)

Theristicus caudatus (Black-faced ,, ). 3 16 4]

T. melanopis (Grey ak) aa 18 31

Plegadis falcinellus (Glossy Lan) ee its) 30:5 112

P. guarauna (White-faced _,, ) 2 es) 4)

Hudocimus, whole genus.................. 43 25°5 143

EL. albus (amity) LSS) Sou coekoe 4) 69°5 143

E. longirostris (Red-billed ,, ) ......... 2 8°5 12

E.. ruber (Scarlet By) etree 36 20 90

Mr. J. H. Gurney (loc. cit. p. 38) records from our own Gardens

a Sacred Ibis alive at 20 years old, and another bird of the same

species from Rotterdam of 11 years. In the list given above

there is a Straw-necked Ibis with a duration of 25 years, and an

Australian Sacred Ibis of over 18 years, whilst there are a number

of other high records. The figures of average duration are very

varied, but in several cases they extend to a number of years, the

low figures being chiefly birds of which there were very few

examples.

In the period in question nearly all these birds had free access

to the open air, but in some cases had shelter in a heated aviary

in winter. Ibises have a simply disposed but rather long in-

testinal tract, with the ceca and the hind-gut much reduced.

It must be inferred from the figures that they are birds of

good viability in proportion to their size.

Summary of Herodiones.

Herons, Storks, Spoonbills and Ibises form a coherent zoolo-

gical group all chiefly carnivorous or piscivorous, with relatively

long, narrow, intestinal tracts, the posterior region of which is

reduced and uneapacious. Within the different groups there is

a fair correspondence between duration and size, but this does

not reach to the whole assemblage, as the Spoonbills and Ibises

show a better viability than the Storks, whilst the Herons are

lowest of all. I see no reason for assuming much difference

(except perhaps in proportion to size) between the potential

longevities of the three divisions, and must assume that the

RELATIVE VIABILITY IN MAMMALS AND BIRDS. Eyl

conditions of captivity have pressed most heavily on the Herons.

I have not sufficient information to point to any special unfavour-

able circumstance in the case of the Herons.

Capt. Flower records no example of the Herodiones still alive

after 12 years.

ODONTOGLOSS A.

PHONICOPTERIDE. Tdi jo isuonth. Soar te

Phenicopterus, whole genus ............ 23 82 273

P. ignipalliatus (South American 5 42 105

Flamingo).

P. roseus (Kuropean Flamingo)...... 9 163 273

P. rubra (Ruddy + Me sesteae 9 21 147

I can find no published statements regarding the longevity of

these large birds, and the maximum duration of nearly 23 years

for a European Flamingo is therefore specially interesting. I

know of no reason for the greater success of the EKurepean form.

The distribution of individual durations with regard to the

maximum durations shows that the deaths with low duration

have nearly always taken place soon after arrival: if the birds

of any of the three species survived two or three months, they lived

years. These birds live on a mixed diet, in captivity taking a

good deal of grain; the alimentary tract is relatively long and

slender, but the posterior portion and the ceca are more capacious

than in most birds. They have been kept with complete exposure

to the open air in summer, whilst in winter, in the period to

which the records refer, they were usually transferred to an aviary

with open-air cages and heated retreats. Flamingoes have a

high longevity and viability under such conditions.

Yo. v. dur. Max. dur.

PALAMEDE A, Toi. es erate in months

Chauna cristata (Crested Screamer)... 11 45°8 115

C'. derbiana (Derbian et Rosa 1 8 16°5 84

Palamedea cornuta (Horned ,, +)... 3 31-1 43

I am unaware of published records as to the longevity of

Screamers. The maximum duration, over 9 years in the case of

a Crested Screamer, is surpassed by two examples of the species,

at present alive in the Gardens, after a duration of nearly 13 years

As in the case of the Flamingoes, the incidence of the mortality

was especially heavy on new arrivals: if the birds lived a few

months, they usually approached the maximum duration recorded.

Screamers are large, bulky birds, with the alimentary tract of a

primitive type, the hind-gut and ceca being relatively capacious.

They have been kept under conditions relatively similar to those

of the Flamingoes, and certainly in proportion to their size

cannot be regarded as highly viable.

512 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

ANSERES.

Cyanine. Tndig |? ganipedta oe

Cygnus, whole genus .........2........ 88 33°09 WoT

C. atratus (Black Swan)... 13 18. 72

C. buccinator. (irumpeter 7) ee.) i DD 144

C. bewicki (Bewick’s Sie ecm: hcg | OS 114

C’. musicus (W hooper Whee sn tues 55°5 132

C’. olor (Mute a Nee. habe A1‘5 WOT

C. nigricollis (Black-necked ,, )... 41 24. OS elenl

Coscoroba candida (Coscoroba ,, )... 7 36 103

Mr. J. H. Gurney has recorded (loc. cit. p. 39), on the authority

of W. J. Broderip,a Mute Swan of 70 years; the highest figure

on my list is not quite 15 years. The distribution of the

individual durations about the average, shows that the heaviest

mortality was in those examples which lived only a few months,

and were either new arrivals, or cygnets; if individuals survived

the early period of detention, they approached the maximum.

PASERANSTINE: Tula) | dafuontli. quan

Anseranas semipalmata (Black and 6 alg 229

White Goose).

PLECTROPTERIN.

Plectropterus, whole genus............... 15 72 152

P. gambensis (Spur-winged Goose) ... 12 82 152

P. niger (Black ,, “ En) Gr hel, Wao 28°5 31

ae rueppellt (Riippell’s ,, ad Va) Wie. Sel 36 36

Cairina moschata (Muscovy Duck) .. 250 9 93

Sarcidiornis carunculata (American 5 33 ley

Black-backed Goose).

S. melanonota (Black-backed Goose)... 8 61 110

hodonessa caryopha yllacea (Pin- -headed 5 16 23

Duck)

Aex galericulata (Mandarin cape eye nex) 42 135

Aex sponsa (Summer as 34 IIL

CEREOPSINE.

Cereopsis novee-hollandie (Cereopsis 13 36 1g)

Goose).

ANSERINA.

(ONoan, IMO) ESN) hoo ec abe soocbEoeaobes- 10 iG 198

C. hyperboreus (Wavy Snow-Goose).. 3 85 92

C’. cerulescens (Blue i a ye Mimics 57 103

C. nivalis (Lesser ,, SSO gee Nis 198

MUOSOP HOM) ETAT) aocoeconssacssonwoe. 41 (i 289

A. albifrons (White-fronted Goose) .. 6 74 168

A. brachyrhynchus (Pink-footed Goose) 4 52 99

A. cinereus (Grey-Lag Goose)...... 12 41:5 126

A. cygnoides (Chinese Sean i 70 153

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

ANSERIN® (cont.). Boia

Anser erythropus (Little Goose)...... 2

A.indicus (Bar-headed ,, )...... 8

A. segetum (Bean iy errr ce: 2

Bernicla, whole genus ....... ean 61

B. brenta (Brent Goose)...... 32

. canadensis (Canada aS Pere 14

BL. hutchinsi (Wutchins’ sey) Fe SHoe 2

4. leucopsis (Bernicle sy yeacierte 12

B. ruficollis (Red-breasted ,,_ )...... 1

Nesochen sandvicensis (Sandwich 9

Island Goose)

Chloephaga, whole genus .............+- 120

C. magellanica (Upland Goose) 83

C. melanoptera (Andean Pear) ee

C. poliocephala (Ashy-headed eS

C. rubidiceps (Ruddy-headed ,, ) 22

Chenonetta jubata (Maned sf) eal

Ay. dur.

in months.

513

Max. dur.

in months.

Geese are popularly reputed to have a very high potential

longevity; Mr. J. H. Gurney (Joc. cit. p. 39) mentions, on the

somewhat doubtful authority of Willughby, a domestic Goose,

80 years old, and on more exact authority a Cereopsis of 33 years

and a Bernicle Goose of 32 years. The highest figure on my list

is 24 years for a Little Goose; but the maximum and average

durations (the latter in many cases are lowered by the inclusions

of broods of goslings) show that these birds have a high viability.

ANATIN. Moyes

Indiv.

Dendrocygna, whole genus............... 99

D. arborea (Black-billed Tree-duck)... 6

D.arcuata (Wandering ,,_ ,, )... 14

D. autumnalis (Red-billed ,, ahs) 32

D. eytont (Kyton’s ,, St A) ici i

D. fulva (Fulvous _,, sti)

D. javanica (Indian he i Seva) 8

D. major (Larger Rn eee ie it

D. widuata (White-faced ,, ,, )... 21

Chenalopex egyptiaca (Egyptian Goose) 16

C. jubatus (Orinoco Goose) ............ 14

Tadorna, whole genus ...............45 109

T. casarca (Ruddy Sheldrake) ...... 22

Ff. cornuta (Common = Nisaetss 31

T. tadornoides (Australian ,, Diaseex: 15

T. variegata (Variegated ,, +)... 4]

Anas; whole; sens), J.1.sj9.s:.woatee-. soon: 117

A. boscas (Wild Duck). = 19

A. cristata (Crested Be ee ee

A. obscura (Dusky pete tt

A. pecilorhyncha (Spot-billed ,, ) 37

Avy. dur.

in months.

Max. dur.

in months.

514 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

' ANATINE (cont.). No. of Ay. dur. Max. dur.

Indiv. in months. in months.

Anas specularis (White-marked Duck) 1 Sih 11

A. superciliosa (Australian Wild ,, ) 45 A2 190

A. undulata (Yellow-billed Sin Falak) 85 319

Chaulelasmus streperus (Gadwall) :.. 10 505 122

Mareca penelope (Wigeon) Ans eect Al 59°5 155

M. sibilatrix (Chiloe Wigeon) ......... 37 A8 185

Dagar acuta) Wpaimtanl\ pe... ene ner 21 5D 214

D. spinicauda (Chilian Pintail)......... 67 40-5 273

Pecilonetta bahamensis (Bahama Duck) 24 DOT 170

P. erythrorhyncha (Red-billed ,, ) 1 128 128

Nesonetta aucklandica (Auckland 1 Ik@f Ii

Island Duck)

Querquedula, whole genus............... 192 AD 208

Q. brasiliensis (Brazilian Teal) ......... 34 39 126

Q. castanea (Chestnut-breasted Duck) 11 19 96

@. circia (Garganey Duck)............... 43 45 164

(). crecea (Common' Peall).---.5.-- 43 52 152

Q. cyanoptera (Blue-winged ,, )......... 5) 13°6 43

Q. falcata (Falcated ca eebige tee 9 18 50

Q). flavirostris (Chilian aa eee tee 8 By 87

@. formosa (Japanese ay peAaNon ae 21 48 150

Q. gibberifrons (Slender Duek) Co areeee 18 75 208

Spatula clypeata (Shoveller) ............ 18 60 154

Marmaronetta angustirosiris (Marbled 12 18 103

: Duck)

FULIGULINA,

Metopiana peposaca (Rosy-billed Duck) 39 AT 160

Puligula, whole genus ..............---. 82 42:5 19s:

ff, baeri (Baers aac.) ie reenan 5 21 AT

10%, fe: EOOTIEC))) . docisoosaddaceda sesso 21 60 193

F. ferruginea (White-eyed Duck)...... 1 il 1

Jal GiOcHrales (CSIGANEID)) - Scoccannendinatiaossdooné 8 20 73

#, rufina (Red-crested Pochard) ...... 24 39 144

Tachyeres cinereus (Steamer Duck) . 1 15 15

Clangula glaucion (Golden-eye) ...... 20 33 164

Gidemia nigra (Common Scoter) ...... 1 30 30

Somateria mollissima (Eider Duck)... 3 14 17

ERISMATURINA.

Biziura lobata (Musk Duck)............ 2 2:5 3

MERGANETTINA.

Hymenolemus malacorhynchus (Soft- 4 3D 72

billed Duck)

MERGINA.

Mergus albellus (Smew) ...............05. 9 22 A6

M. merganser (Goosander)............... 5 35 84

M. serrator (Red-breasted Merganser). 3 7 18

RELATIVE VIABILITY IN MAMMALS AND BIRDS, 515

Mr. J. H. Gurney gives (loc. cit. p. 39) one doubtful case of a

Wild Duck of 29 years, and several well authenticated examples

of various ducks ranging from 23 years downwards. ‘The figures

of maximum duration on my list are almost uniformly high, the

best being a Yellow-billed Duck of 26 years’, a Chilian Pintail

of nearly 23 years’, and a Red-billed Tree-duck of over 20 years’

duration. The figures of average duration are reduced in many

cases by the inclusion of a number of very short-lived birds, either

ducklings or new arrivals, but none the less show a fairly consistent

high viability.

Summary of Anseres.

The first question that comes into special prominence in the

case of Swans, Geese and Ducks is the validity of the units on

which these averages are made. The individuality of these birds

is difficult to distinguish, and as in the period to which the

figures relate, there was no system of marking the individuals,

mistakes must have been easy to make, where opportunity

occurred. The swans, geese, and a good many of the larger

ducks, such as Sheldrakes, were kept in separate pens, and the

chances of confusing individuals were very small. In the case of

birds like the Mandarins, Summer Ducks, Wild Duck, Pin-

tail, Teal, and so forth, a good many were kept in the same

ponds, and mistakes as to individuals must have taken place.

However, the numbers at any particular time during the whole

period of 33 years were never very large; and if the figures of

cases where individuals were very few and were kept separate be

compared with those where confusion was more likely, the

inference can be drawn fairly, that on the whole such mistakes

have corrected each other, and that the results are fairly valid.

It is to be noticed that the familiar correlation between size

and longevity does not appear in these figures of Anserine birds,

although it may apply to potential longevity. There is very

little to choose between the ducks and geese, and both ducks and

geese on the whole show higher maximum and average durations

than occur for swans. In all the Anseres the alimentary canal is

relatively long, arranged in a somewhat complex fashion, and

narrow in calibre; the hind-gut and ceca are, in comparison with

most other birds, rather long, and capacious (except in the

Mergansers, where the ceca are vestigial). They are highly

intelligent birds, readily adapting themselves to new conditions

and realizing quickly that they have nothing to fear from man.

Except that the space given to Swans has been rather limited,

the conditions have been good, as these birds, whether from the

tropics or not, have been invariably given access to open air, and

in most cases have spent their whole time in it. The figures of

average and maximum duration show that these birds havea good

viability in captivity. None the less, as the conditions are rather

unusually favourable, I think the inference can be drawn that, in

516 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

proportion to their size, Anseres have not naturally very high

longevity or viability.

Capt. Flower records only one example of the Anseres—an

Egyptian Goose—still alive after 12 years.

COLUMBAL.

TRERONIDE. TOe, Ghacalin Bao le

Sphenocercus sphenurus (Wedge-tailed 1 35 35

Pigeon).

Vinago. whole tsenus eee eens eee C 6-4 9

V. calva (Bare- faced Fruit- ee) 5 6 5)

V. crassirostris (Thick-billed ,, ) ... 1 4 4

V. delalandiit (Delalande’s ,, ) . 1 9 9

Crocopus chlorogaster (Southern mt 34 34

; Fruit-Pigeon)

C. phenicopterus (Purple-shouldered 1 a) 5

Fruit- Pigeon).

_ Osmotreron bicincta (Double-banded = 1 4 4

Fruit-Pigeon).

O. vernans (Parrot : B 2 10 14

Ptilopus, whole genus:...1....2..........- 9 DD 22

P. fasciatus (Banded Fruit-Pigeon)... 2 1 1

P.jambu (Jambu ,, a eae eae 6°5 22

P. melanocephalus (Black-headed 2 10 11

Fruit-Pigeon).

P. porphyraceus (Purple-crowned i 1 1

Fruit-Pigeon).

Globicera, whole genus .................. 5 24 65

G. aurore (Aurora Island Fruit-Pigeon) 1 65 65

G. pacifica (Pacific 2, 8°5 9

G. rufigula (Vinous-thr oated Fruit: 2 18:5 26

Pigeon).

Carpophagacenea(Bronze Fruit-Pigeon) 10 Mare (| 53

C. paulina (Red-naped ,, nA 4 27 4]

Myristicivora luctuosa (Nutmeg Fruit- 4 14 16

Pigeon).

Lophemus antarcticus (Double-crested 17 13 25

Fruit-Pigeon).

_COLUMBID&.

Columba, whole genus ................5. 138 34-7 227

C. albigularis (White-throated Violet 1 g) 9

Pigeon).

C. araucana (Araucanian ee eas! ag 25 25

C. bellit (Bolle’s as 6 22 65

C. caribbea (Ring-tailed sy ul 56 56

C. eversmannt (Kversmann’s 5

C. guinea (Triangular-spotted _,,

C. gymnophthalma (Naked-eyed_,,

O. inornata (Cuban “8

YS SS

for)

eo)

(or)

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

CoLuMBID# (cont.).

Columba laurivora (Canary Pigeon)

C. leucocephala( White-crowned ,, )

('. leuwconota (White-backed £ Wa)

C’. maculosa (Spotted sal)

C'. cenas (Stock-Dove) ............

C. palumbus (Ring-Dove) ............

C’. picazuro (Picazuro Pigeon)...

C. plumbea ( Vinaceous ati h INS J

C. rufina (Rufous Ty ee

C'. speciosa (Specious 5

Gs squamosa (Porto Rico ae ce

Macropygia leptogrammica (Narrow-

barred Pigeon).

M. phasianella (Pheasant-tailed ,, )

Eectopistes migratorius ( Passenger

Pigeon).

Zenudura carolinensis (Carolina Dove)

Fenaida, whole senus .,.... 2.5. .in-...e.

ah Pee (Zenaida Dove) Serene:

Z. auriculata (Auriculated Dove)......

Z. wurita (Martimscam =e) -..2:

Nesopelia galapagoensis (Galapagan

Dove).

Melopelia leucoptera (White-winged

Zenaida Dove).

T. aldabranus (Aldabra Turtle Dove)

T. bitorquatus (Double-ringed Turtle

Dove).

T. capicola (Cape Turtle Dove)

T. chinensis (Chinese _,, te)

T. communis (Turtle Dove) .........

T. orientalis (Eastern Turtle Dove)

T. risorius (Barbary ,,

T. semitorquatus (Half-collared Turtle

Dove),

7. senegalensis(Cambayan Turtle Dove)

T’. suratensis (Spotted a pe)

T. tigrinus (Tigrine a ean)

T. vinaceus (Vinaceous _,, ew)

Geopelia, whole genus.....................

G. cuneata (Graceful Ground-Dove) ...

G. humeralis (Barred-shouldered Dove)

G. maugei (Maugeé’s Dove)............

G. striata (Barred Fae) AS ee Or

G. tranquilla (Peaceful ,, )............

Scardafella squamosa (Sealy Ground-

Dove).

Columbula picui (Picui Dove) .........

No. of

Indiv.

ee ty —

SE OLD Op

— me bd

DCH H OS

Proc. Zoou. Soc.— 1911, No. XXXIV.

Avy. dur.

in montl

40)

33°5

1s.

138

D17

Max. dur.

in months.

aire

100

ail

166

215

518 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

CoLUMBIDH (cont.). Te ee ae oie

Chamepelia passerina (Passerine 95 11:8 27

Ground- Dove).

C. talpacoti (Talpacoti < ) 16 30 67

Peristera geoffroyi (Geoffroy’s Dove) 34 26°5 102

Metriopelia melanoptera (Black-winged 8 29°5 121

Dove).

(na capensis (Cape Dove) ............ 1] 28 69

Tympanistria bicolor (Tambourme 16 26 61

Pigeon).

Chalcopelia chalcospilos (Bronze-spotted 17 By ats) 92

Dove).

Chalcophaps chrysochlora (Little Green- 5 32 61

winged Dove).

CO. indica (Green-winged Dove)......... 23 3D 193

Calopelia puella (Schlegel’s ,, )......... 18 21 109

JEU KCFIS, WINCIS EAMES Sonsceoodassooscseso% 38 24°5 98

P. chalcoptera (Bronze-winged Pigeon) 34 24 98

ie elegans (Brush Bs A Wiercic) 20 28

P. histrionica (Har lequin -Bronze- 1 60 60

winged Pigeon).

Geophaps plumifera (Plumed Ground- 9 26 66

Dove).

G. scripte (Partridge Bronze-winged 5 16 24

Pigeon).

Ocyphaps lophotes (Crested Pigeon) ... 49 22 64

Haplopetia larvata (Riufous-necked 2 14 16

Wood-Dove).

Leptoptila, whole genus ..................

L. jamaicensis (White-fronted Dove)

L. ochroptera (Orange-winged ,, ) 2 iat 14

L. rufaxilla (Red-ander-winged ,, ) 9 25 81

Geotrygon, whole genus ...............-.. 22 27 56

&. cristata (Mountain-witeh Ground- 4 17 25

Dove).

&. montana (Red Ground-Dove) ...... 14 30 56

&. mystacea (Moustache Ground-Dove) 4 29 d4

Phlogenas luzonica (Blood-breasted 29 24 79

Pigeon).

P. stair (Stair’s Ground-Dove)......... ZI 14°5 18

Leucosarcia picata (Wonga-Wonga 22 50 171

Pigeon).

Starneenas cyanecephale (Blue-headed 13 20 44

Pigeon).

Calenas nicobarica (Nicobar | ,, ) 23 51 109

Goura coronata (Common Crowned 30 23 110

Pigeon).

@. victorice (Victoria Crowned _,, 1) 58 142

Didunculus strigirestris (Toothed- 3 30 53

billed Pigeon).

RELATIVE VIABILITY IN MAMMALS AND BIRDS. SS)

Summary of Columbe.

Mr. J. H. Gurney (loc. cit. p. 39) gives on the authority of

Mr. F, E. Blaauw several instances of long-lived Collared Doves

(Turtur risorius), ranging from 40 to 30 years, and a Common

Pigeon of 284 years; a correspondent of my own, Mr. E. Mellin,

has informed me of a Ring-Dove of over 23 years old. The

figures of maximum duration on my list are therefore not

surprising: they include a Vinaceous Pigeon of just under 19

years, a Porto Rico Pigeon of just under 18 years, an Eastern

Turtle-Dove and a Green-winged Dove of over 16 years. The

maximum durations are on the whole good ; certainly in proportion

to size better than inthe Anseres. The fleur és of average duration

are much lower in the Fruit Pigeons, in w which they seldom exceed

2 years and are frequently under | year, than in the Columbide,

where they mostly range from 2 to 5 years. J cannot see any

good correlation within the group, between size and viability.

Pigeons have a long and complex alimentary tract, with,

however, the posterior portion and cca relatively much reduced,

They have been kept under various conditions in the period under

discussion, but most of the tropical forms have been placed in

a heated aviary, sometimes with, sometimes without access to the

open air.

Capt. Flower’s highest record is that of a Green-winged Dove,

alive after 6 years’ duration.

dC ‘ No. of Av. dur. Max. dur.

PTEROCLETES. ees Av. dur. Max. dur

in months. in months.

Pierocles, whole genus...................+5 67 20°5 94

P. alchata (Pin-tailed Sand-grouse) ... 18 15 34

P. arenarius (Black-belliel Sand- 27 21 55

grouse).

P. bicinctus (Double-banded Sand- 9 17 32

grouse).

P. exustus (Little Pin-tailed Sand- 11 31 94

grouse).

P. fasciatus (Banded Sand-grouse) 1 24 24 —

P.namagua (Namaqua_,, Jat wall 15 15

Syrrhaptes paradorus (Pallas’ Band- 16 8 14

grouse).

I had no previous information as to the longevity of Sand-

grouse. The maximum durations, of which the highest is under

8 years, and the average durations, most of which are considerably

less than 2 years, show that these birds, in proportion to their size,

are less viable than Dovesand Pigeons. Their alimentary tract is

somewhat similar to that of Pigeons, but the hind-gut and ceca

are relatively capacious. In the period under consideration Sand-

grouse were placed in an aviary with heated interior and open-air

runs.

Capt. Flower records Spotted Sand-grouse (P. senegallus) still

alive after over 9 years’ duration; a year longer than the highest

record I give.

34%

520 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

GALLIN A.

Tap vo. of

TETRAONID&. ee

Uetrgonwinole yo ei Siro rere eect ee ee ne 11

T, cupido (Praimie Grouse)... -.2c:----.7- 7

T. phasianellus (Sharp-tailed Grouse). 2

T. wrogallus (Capercaillie) ............... 2

PHASIANIDA,

Galloperdi 79, WOES EXETMINS 5 soe Secncooun 5

7. lunulata (Hardwicke’ s Spur-fowl) . t

G. spadicea (Rufous os sf 2

G. zeylonensis (Ceylonese 5 5) Rene

ithaginis geoffroyi (Geoftroy’s Blood- 1

Pheasant).

Ptilopachys ventralis (Buff-breasted 2

Partridge).

Francolinus, whole genus ............... 68

F. bicalearatus (Double-spurred 7

Francolin).

F. capensis (Cape Fr. ancolin) Kl eens 1

FP. clappertont (Clapperton’s Francolin). 8

EF. coqui (Coqui iti) asl

Lf. granta (Grant's oy Me nee

I’, gularis (Wood A Need

FP. kirki (Kirk’s fe

F, levaillante (levaillants ,, ). 1

F. natalensis (Natal i poma Tae]

F. pictus (Painted gece Ola

F. ponticerianus (Grey osm) say ee)

F, vulgaris (Black. va) earls

Pternistes, whole genus .................- 29

P. afer (Grey- winged Francolin) 7

P. infuscatus ( Dusky Hf yee a

P. leucoscepus (White-shafted | ,, aa aT:

P. nudicollis (Bare-necked - eae

P., swainsont (Swainson’ S aed,

Arboricola atrogularis (Black- Aiveated 1

Francolin).

A. torqueola (Hill Francolin)............ _)

Bambusicola fytchii(Fytch’s Francolin). — 1

B. thoracica (Bamboo Partridge) ...... 15

Perdi« cinerea (Partridge) ns ee SR 14

P. hodgsonice (Eodgson’s Partridge)... 4

Coturnix, whole genus .................. 108

C. aineons ‘(Chinese @iranl) eee US}

C. communis (Commionaie i) eee 54

O. coromandelica (Rain ka NS. ea Is

C. histrionica (Harlequin ,, )...... 12

C. pectoralis (Bectoral yar) eer 4

Av. dur.

in months,

9-7

8:5

Max. dur.

in months.

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

PHASIANIDE (cont.).

Synecus australis (Austvalian ine

Perdicula argoondah (Marbled i)

P. asiatica (Asiatic 7)

Rollulus cristatus (Crowned 33)

Odontophorus dentatus (Capoeira ,,_)

O. guianensis (Guiana. )

Ortyx virgimanus (Virginian Colin)

Eupsychortyx cristatus (Crested ,,_)

E, leylandii (Leyland’s Colin) .........

Callipepla, whole genus................45

C. californica (Californian Quail) ......

C. gambelli (Gambel’s es

C. squamata (Mexican ee

Caccabis, whole genus ......0....c05.00

C. chukar (Chukar Partridge) ..:......

C. Ta ee (Black-headed Part-

~——~

ridge).

C. petrosa (Barbary Partridge) .........

C. rufa (Red-legged Niclas oj)ipeeicitesrsses

C. saxatilis (Greek Jace yal Rencttdetoree

Ammoperdix bonhami (Bonham’s

Partridge).

Meneye (levis barhridge) .:' (ees sy..

Tetraogallus himalayensis (Himalayan

Snow Partridge).

‘Lophophorus, whole genus...............

L. impeyanus (Himalayan Monaul) ...

L. Uhuysi (Lhuys’s EN ten:

LI. sclateri (Sclater’s a

Crossoptilon mantchuricum (Ma ntchu-

rian Crossuptilon)

C. tibetanwm (Tibetan ‘5 )

Phasianus, whole genus................5.

P. colchicus (Common Pheasant)

P. decollatus (Collarless ke

P. ellioti (Elliot's 3

P. principalis (Prince of Wales's ,,

P. reevesi (Bar-tailed i

P. semmerringi (Semmerring’s ,,

P. torquatus (Ring-necked ,,

P. versicolor (Japanese x

P. wallichia (Cheer

Thaumalea amherstice (Lady Amherst’s

Pheasant)

T. picta (Gold Pheasant) ...............

Pucras sia, WIHOLCL SoTL Revs. entree ere

P. darwint (Dar win’s Pucras)

P. macrolopha (Indian m

P. xanthospila (Yellow-spotted ,, _ )

No. of

Indiv.

Ay. dur. Max. dur.

in months. in months.

26°7 57

28 28

12°4 27

9 20

9:9 29

74 U¢/

il 38

14 58

43 43

18 9]

a 9]

26 49

18 46

12 50

11 50

9 20

1133 50

SG) 31

15 32

20 4]

5) 19

3 a

236 123

23 123

54 04

20 20

22°8 49

22, 32

DOO 95

26 85

18 18

oe 76

18 46

22°8 64

36 95

20 70

20 86

31 79

20 97

22°5 93

8°6 24.

9 24

3 3

4 4

522 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

PHASIANIDE (cont.). Twiiy, cana Peat:

Euplocamus, whole genus ...........7... 155 24°8 161

E. albo-cristatus (W hite-crested Kaleege) 12 36 76

E. andersoni (Anderson’s Bites) yn 101 ll

E. erythrophthalmus (Rufous-tailed 18 20 109

Pheasant)

E. horsfieldi (Purple Kaleege) 7 46 108

EL. lineatws (Lineated sia ORO 22 76

EH. melanotus (Black-headed ,, ) 8 45 16]

EL. nycthemerus (Silver Pheasant)...... 38 23 114

E.. nobilis (iiige= laren) ere r 8 37 116

EL. prelatus (Siamese ue ann eae 8 40 1h

He swinhow VW iSwinhoe’s) 5)... 32 12 66

&, vieilloti (Vieillot’s Fire-back) ...... 4 11 21

Gallusmwholeycenus-reete toe sane 39 22°7 148

G. bankiva (Bankiva Jungle-fowl)... 5 8 36

G. sonnerati (Sonnerat’s __,, iS) 12 ou 148

G. stanleyi (Ceylonese ,, a) ipl 18 36

G. varius (Fork-tailed _,, Pe casi paltlt Wes 62

Ceriornis, wihtole Sens \(.:.-)--.5.--.---- a 29 133

C. blytht (Blyth’s Tragopan) ............ 5 38 133

C. caboti (Cabot’s mh ) cia cals eae 6 31 le

C. melanocephala (Black-headed 2 26°5 28

Tragopan)

C. satyra (Horned Ad yr 2h ingle?) 58

C. temminckt (Temminck’s _,, ar alt 30°5 123

JEM. WANE, GENUS sadcboodsssoosboopsoncoe's 99 22°6 98

P. cristatus (Common Peafowl) 66 24 98

P. nigripennis (Black-winged . ,, ) 15 26 73

P. spicifer (Javan a} agellts) 1133 47

Polyplectron, whole genus ............... AT 50 179

P. bicalearatum (Crested Peacock 2 23 4]

Pheasant)

P. chinquis (Peacock Pheasant) ...... 39 55 179

P. germaint (Germain’s Peacock 6 22 AT

Pheasant).

Argus giganteus (Argus Pheasant). 22 42 163

Meleagris gallopavo (North American 12 26 62

Turkey).

M. ocellata (Ocellated Turkey) ......... 2 2°5 3

Nunuda, wholegenus ~.-..2............ 34 aa 40

NV. ellioti (Elliot’s Guinea-fow] 1 4) 5

N.meleagris (Common ,, a 8 28 40

N. ptilorhyncha (Abyssinian

N. reichenowi (Reichenow’ s

77 ole)

NV. nutrata (Matired a aid) all 5 30

)

>) ?

G. cristata (Crested Guinea-fowl)

G. edouardt (Verreaux’s ,, CN sh atl! rag Ie 21

RELATIVE VIABILITY IN MAMMALS AND BIRDS,

PHASIANIDE (cont.).

Guttera pucherani (Pucheran’s Guinea-

fowl).

Acryllium — vulturinum (Vulturine

Guinea-fow]).

CRrAcID a.

O70, MUWOLe’ SOWUS 2a nce. ee ce ce cow onaie

C. alberti (Prince Albert’s Curassow)

”, alector (Crested or)

C. carunculata (Yarrell’s =

C. daubentont (Daubenton’s rd)

". globicera (Globose shores)

C. globidosa — (Globulose Ay) Pics)

C. incommoda (Inconvenient singh)

C. sclateri (Sclater’s tea)

Mitua tomentosa (Lesser Razor-billed

Curassow).

M.tuberosa (Razor-billed ,, )

Pausis galeata (Galeated ‘5

Nothocrax wurumutum (Urumutum

Curassow).

Penelope, whole genus ..................

P. cristata (Rufous-vented Guan)

P. greeyt (Greey’s -

P. jacucaca (White-fronted 5

P. marail (Marail .

P. pileata (Red-breasted ’

P. purpurascens (Purplish ss

P. superciliaris (White-eyebrowed ,,

mooie. WOle SEMWUS ya.c-..2-c.cece se. -ls0

P. cujubit (Amazonian Guan) .........

P. cumanensis (Piping Guan)

P. jacutinga (White-crested _,,

Alburria carunculata (Wattled ,,_ )

Oniniiss WHOVG CNUs ors. 5..001 ec: «tenes

O. albiventris (White-bellied Guan)

)

O. garrula (Chattering sees)

O. motmot (Little ue)

O. ruficauda (Red-tailed en)

O. vetula (Mexican re)

MuEGAPODIIDA.

Talegalla lathami (Brush-Turkey)......

Megacephalon maleo (Great-headed

: Maleo).

Leipou ocellata (Mallee Bird)............

No. of

Indiv.

Dmriwewesatbnwe

H oo HS = Go «

Summary of Galline.

Mr. J. H. Gurney (loc. cit. p. 39) gives a few interesting figures

for Gallinaceous birds, including domestic fowls of 30, 25 and

24 years, and a Silver Pheasant of 21 years.

Ay. dur,

in months.

14-4

523

Max. dur.

in months.

101

The maximum

524 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

durations on my list include some fairly high figures—a Brush-

Turkey of over 19 years, a Peacock Pheasant of nearly 15 years,

Argus and Kaleege Pheasants over 13 years, a Monaul of over 10

years and a Common Quail of over 7 years. Taking Galline as a

whole there is no close correlation between size and duration.

The Grouse are the least viable in captivity, the average durations

being nearly all under a year, and 4 years being the maximum.

The Francolins, Quails, and Partridges have an average duration

most often between 1 and 2 years, with a few good figures of

maximum duration. The Pheasants, Peacocks, and Jungle-Fowl

for the most part have average durations exceeding 2 years, and

reach 13 to 15 years maximum. Sir H. G. lhavien Wisks VALS.

however, has told me of a Peacock still alive at a house near

Aylesbury, and which is certainly 40 years old. The Curassows

and Guans appear to be specially viable; there is a remarkable

uniformity amongst the individual durations, these lying very

close to the average, which is generally over 3 years.

The strueture of the intestinal tract does not differ much in

Gallinaceous birds: in all the gut is relatively capacious, although

not specially long, and the posterior portion including the czeca is

long and capacious compared to the whole gut. It is perhaps

not more than a coincidence that the hind-gut and ceca are rela-

tively least capacious in the Megapodes, ne least capacious in the

Curassows, and most capacious in the other Gallinaceous birds,

whilst the Megapodes show the best maximum duration, and the

Curassows the highest average durations in the group. The

conditions under which the birds of this group were kept during

the period i in question did not differ much; all had access to open

air, and the smaller Quails, Partridges and so forth, and the

Cur. assows had warmed shelters, whilst the others ‘hel unheated

sheds. Considering the more or less favourable conditions, and

the large size of many of the Gallinaceous birds, it cannot be said

that they display a good constitutional viability.

Capt. Flower’s records are closely comparable with the maxima

I give here. He notes a Clapperton’s Francolin still alive after

6 years, my highest record being 5 years; a Quail of over 7 years,

the same maximum as mine; an Amherst Pheasant of 11 years’

duration, and a Silver Pheasant still alive after 12 years; these

are higher than the records of the same species I quote, but, on

the other hand, I cite a Black-headed Kaleege of 13 years piiighal

Peacock Pheasant of 15 years.

Yo. of Av. dur. x. dur.

HEMIPODII. ie mn Rene incon

Turn, whole genus <.2,.......6.-.40015. 15 9 26

7’. dussumieri (Little Bittern Quail)... 2 2°5 4

7’. lepurana (Lepurana Hemipode)... 3 AG 26

1’. sykest — (Sykes’s - Nasa ate 8 16

7’. taigoor (Black-breasted , Veneer nee 1 1

7. tanki — (Tipperah BE ve 2 1 l

1. varia (Varied 4% yew 4 16 29

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 529

T am ignorant of any figures hitherto recorded as to the

longevity of Hemipodes. My figures show a very low viability

both from the point of view of average and maximum dura-

tions. The birds were kept during the period dealt with under

the same conditions as the small Francolins, Quails and

Partridges. The alimentary tract is relatively short but

capacious, the hind-gut and ceca being very capacious relatively

to the size of the birds. The group has a low viability.

FULICARLA.

RaLuip2. Thuliv, a aa “= ai an :

mates, whole genus’ .....66)...0eeceebees 24 Pes 5]

Rh. aquaticus (Water-Rail)............... 10 13-9 35

R. celebensis (Celebean Rail) ............ 3 36 48

R. maculatus (White-spotted Rail) ... 3 26 48

R. pectoralis (Australian of) 6 19°5 35

R. rhytirhynchus (Sooty Sse mee 36 51

Aramides cayennensis (West-Indiai 19 10 26

Rail)

A. ypecaha (Ypecaha Rail) ............ Ih 15°5 39

POTLONG, WHOLE SONUS 60... yee geen: 5 30 59

P. carolina (Carolina Crake) ......... 2 22°5 28

P. maruetta (Spotted ,, ).........65. 2, 29 29

Haanorana “(Wiariked “4 \l i ecc veces. 1

Rallina peciloptera (Bar-winged Rail) 2 2

Crex pratensis (Corn Crake) ............ 16 22 101

Ocydromus, whole genus ........ ...++. 30 52 ILI

O. australis (Weka Rail) .............+. hi 49 Waly

acai dbialeles. syae eyo!) kere esses: 8 64 107

O. fuscus (Black Woodhen) ............ 1 50 50

O. lafresnayanus (New Caledonian

Rail) 2 A3°5 76

O. sylvestris (Woodhen Rail)..........-. 2 38°5 66

Limnocorax niger (Black Gallinule)... 2 52°5 89

Porphyrio, whole genus.............+.+++ D3 23°5 (5

P. calous (Javan Porphyrio) ...........- ] 1 1

P. ceruleus (Purple Gallinule) ......... 4 21 32

P. edwardsi (Miine Edwards's 2 18°5 22

Porphyrio)

P. madagascariensis (Madagascar 17 18 59

Porphyrio)

P. melanotus — (Black-backed ,, yay ae) 25 73

P. poliocephalus (Grey-headed ,, yes as) 32 75

P. vitiensis (Grey-blue _,, eal 23 23

Hydrornia alleni ( Allen’s a yes: 17 47

Tonornis martinicus (Martinique 8 We 30

Gallinule)

Tritonyx mortiert (Mortier’s Water- 1 102 102

hen).

7. ventralis (Black-tailed Waterhen). 3 85 162

526 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

No. of Ay. dur. Max. dur.

Rauiip (cont.). Indiv. in months. in months.

Gallinula, whole genus ../..........2-++- 20 25°5 42

G. chloropus (Moorhen) .................. 9 30 42

G. galeata (American Moorhen) ...... 2 21 29

G. nesiotis (Island-Hen Gallinule) 3 25 35

G. phenicura (White-breasted ,, ) 4 12°5 26

G. pyrrhorhoa (Madagascar Moorhen) 2 38°9 42

EulC am aWiNOle eens. Yaseen ert e 8 14 36

fF. ardesiaca (Slaty Coot) ............... 2 18°5 36

Wisatral Coop) ive kt ks. neeeree es. Sosa 4 16 21

F. leucoptera (Strickland’s Coot) ...... 2 6 9

Summary of Fulicarie.

I have discovered few published statements as to the longevity

of Rails and Crakes, but Mr. J. H. Gurney (loc. cit. p. 38) has

mentioned Porphyrios of 19 and 14 years. The highest figure

on my list is that for a Black-tailed Waterhen with a duration

of over 13 years; then come a Weka Rail of over 9 years; a

Corn Crake of 8 years, and then figures ranging downwards.

The average durations range rather closely round 2 years. The

intestinal tract of Rails and their allies is of moderate length

end capacity, the posterior portion with the ceca being rather

capacious. These birds in the period in question have been kept

either in open-air paddocks, or in an aviary with heated interior.

and access to open air. There is not a very wide gap between

the figures of average and maximum durations; there is no

appreciable correlation between size and longevity, and considering

the rather favourable conditions under which the birds were kept,

they appear to have a rather low viability in proportion to their

size.

Capt. Flower’s highest record is a Gallinule still alive after

6 years’ duration.

ALECTORIDES.

OTIDIDE. Tae Seca REGGE

Otis tarda (Great Bustard) ............ 16 30 128

Hupodotis, whole genus ....:............. 13 507 106

ZH. arabs (Arabian Bustard)......... 2 D9 91

#. australis( Australian ,,° )......... 4 13 106

Li. caffra (Kaffir ge ee 1 J] 9

LH. denhami (Denham’s Dy fe eae 3 60 98

E. kori (Borchellise yay une) s-ceeers. 2 24 42

E. ruficollis (Red-necked ,, _ )......... 1 10 10

Houbara macqueent (Macqueen’s 8 9 25

Bustard)

H. undulata (Houbara Bustard) ...... 7 3 9

I have found no other figures published as to the longevity of

Bustards. The maximum duration on my list, of nearly 11 years

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 527

fora Great Bustard, is no great age for so large a bird ; the figures of

average duration are fairly good in comparison with the maximum

durations. These birds have been kept almost altogether in open-

air paddocks, and it would appear that their constitutional

viability is not high in proportion to their size. The whole

alimentary tract is very short in proportion to the size of the

birds, but the hind-gut and ceca are relatively extremely

capacious.

Denese ae Cima No. of Ay. dur. Max. dur.

Indiv. in months. in months.

Wesophid, WHOLE GENUS ......:<.0-.s--e0008-+s 30 Wee 90

P. crepitans (Common Trumpeter) ... 15 Wes 90

P. leucoptera (White-backed ,, ) ... 6 19°5 34

P.viridis (Green-winged , ) ... 9 16°5 32

Cariama cristata (Brazilian Cariama) 20 AQ 151

Chunga burmeisteri (Burmeister’s 6 3l 65

Cariama).

IT do not know of any published references to the ages of

Trumpeters and Cariamas. The alimentary tract of these birds

is much of the same character as that of the Bustards, but the

ceca and hind-gut are relatively rather less capacious. The birds

‘have been kept in an aviary with heated interior and open-air

compartments. The record of between 12 and 13 years for a

Brazilian Cariama, and the figures of average duration appear

to show that the Cariamas have a higher viability than the

Trumpeters. ‘The former are larger birds, and the difference

may be merely the familiar size correlation.

No. of Ay. dur. Max. dur.

GRUID&. Indiv. in months. in months.

CUS HWIMOLEISEMUS . 6. 2. cee). Bebe 42 72 514

G. americana (American Crane) ...... 5 34 82

G. antigone (Kastern Sarus ,, ) ...... a 64 191

G. australasiana (Australian Crane) 10 68 129

G. canadensis (Canadian siete) ee 35 96

G. communis (Common an em 145 514

G. japonensis (Mantchurian 1 Mens 59 306

G. monachus (Hooded iain wage 99 99

Anthropoides, whole genus............... A() 46°5 227

A. carunculata (Wattled Crane) ...... 9 34 59

A. leucauchen (White-necked Crane) | 184 184

A. leucogeranos (Asiatic White ,, ) 9 34 84

A. paradisea (Stanley Pe xs, 58 227

A. virgo (Demoiselle ei): sel 46 104

Balearica pavonina (Crowned Pieayt Va 50 141

B. regulorum (Cape Bo psbA 43 170

Mr. J. H. Gurney has recorded (loc. cit. p. 38) a Common Crane

of 40 years (from the Gardens of this Society) and a White-necked

Crane of 28 years from the Amsterdam Gardens. The longest

528 ; DR. P. CHALMERS MITCHELL ON LONGEVITY AND

duration on my list isa Common Crane which lived in the Gardens

for nearly 43 years, another of the same species with a duration

of 274 years, and a Mantchurian Crane of 253 years. It is clear

that these large birds have a very high potential longevity. The

alimentary tract is relatively very long and of narrow calibre,

but the ceca and hind-gut are by no means capacious in proportion

to the whole tract. In the period to which the figures relate,

the Cranes were kept in an open-air paddock, a few being taken

indoors in winter. The maximum durations, and the average

durations, ranging from nearly three years upwards, show that

these birds have a high viability.

No. of Ay. dur. Max. dur.

Hurypycm 2. Indiv. in months. in months.

Kurypyge helias (Sun-bittern) ......... ital 48 114

Rhinochetus jubatus (Kagu) ............ 6 137 196

¥ do not know of any published figures relating to the longevity

of the isolated birds here placed together. The individual examples

of the Sun-bittern differed considerably in duration: all but one

of the six Kagus lived at least 9 years in the Gardens, the

maximum duration being over 16 years. They were kept under

sunilar conditions, in an aviary with warmed retreats and access

to open air. The maximum durations and average durations—

show that both have a high potential longevity and viability in

proportion to their size, but that the Kagu is hardier than the

Sun-bittern. In each case the alimentary tract is, relatively to

the size of the bird, rather long, but the posterior region and

creca are uncapacious.

ARAMID. Noxon Ay. dur. Max. dur.

Indiv. in months. in months.

Aramus scolopaceus (Scolopaceous 1 9 9

Coulan).

IT cannot draw any conclusion from the single example of this

bird. From the structure of the alimentary tr ract, I am disposed

to regard this bird as less viable than Cranes, Bustards, Sun-

bitterns and so forth.

Summary of Alectorides.

Most naturalists would agree that the association of the birds

here grouped as Alectorides (following the Society’s Vertebrate

List) 1s not intimate. It is quite plain that within the assemblage

there is no good correlation between size and longevity and

viability. The Cranes, it is true, show the longest dur ations and

very good averages, but the Kagus, which are Touch smaller, are

nearly as good, whilst the Bustards, which are large and bulky

birds, are much worse. If we make, so to say, a correction for

Size, discounting the expected eveater longevity of the HE er

florins, then the birds in the assemblage might be arranged - as

follows, in the order of good to less good viability Kaus, Crates,

Sun- bitterns. Cariamas, Trumpeters, Bustards generally, and

lowest of all Houbara Bustards.

yan ae

RELATIVE VIABILITY IN MAMMALS AND BIRDS.

LIMICOLAs.

CEDICNEMID.®. Bees

Wdicnemus, whole genus .............6. 31

(1. bistriatus (Double- -striped Thicknee) 4

. grallarius (Australian M3 LS

. scolopax (Stone-Curlew) ............ 1)

(H. superciliaris (Peruvian Thicknee) 8

CHARADRIID&.

Charadrius pluvialis (Golden mi 27

Squatarola helvetica (Grey org 3)

Aigialitis curonica (Little Ringed , DP a2

A, hiaticula (Ringed ssneivaee

Hudromias morinellus (Dotterel) ...... i

Vanellus cayennensis (Cayenne Lapwing) 19

V. cristatus (=vulgaris) (Lapwing) ... 31

Sarciophorus pectoralis (Black-breasted 7

Peewit)

Strepsilas interpres (Turnstone) ...... 6

Hematopus niger (Black Oyster- 2

catcher)

H. ostralegus (Oyster-catcher) ......... 28

Glareola ocularis (Hye-marked = 1

Pratincole)

G. pratincola (Pratincole) ............... 10

CHIONIDE.

Chionis alba (Yellow-billed Sheathbill) 12

C. minor —_ (Black-billed a yy

SCOLOPACID.!.

Recurvirostra avocetta (Avocet).........

Himantopus brasiliensis (Brazilian

Stilt-Plover)

H. nigricollis (Black-necked Stilt)....... 12

Scolopax rusticola (Woodcock) ......... 6

Gallinago colestis (Snipe) .............- 1

Tringa alpine (Dunlin) ........5...0s.6+- DD

CORE US: (ISTOb) 76 isc uep comand doneiues 44

Machetes pugnasx (Ruff) ..........,.2..... 5

Calidris arenaria (Sanderling) ......... ul

Tringoides hypoleucus (Common Sand- 4

piper)

Totanus calidris (Redshank) ............ 19

DS fiscus (SPORLEG: = Fc, AO) Score escecs: 1

Limosa cegocephala (Black-tailed Godwit) 4

L. lapponica (Bar-tailed Godwit) ...... 3

Numenius, whole genus .................. 25

ON. angi (Curlew) 002 co. gace- anaes 20

NV. femoralis (Pacific Curlew)............ 2

NV. pheopus(W bimibrel).. ..d.. nse 6s 3

Ay. dur.

im months.

D29

Max. dur.

in months

WAZ

2a)

530 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

Summary of Limicole.

Mr. J. H. Gurney (loc. cit. p. 40) records an Oyster-catcher of

30 years, and a Ruff of 10 years. The highest figures of

maximum duration on my list are 15 years for a Pacific Curlew

and over 14 years for a Black-breasted Peewit, but there is at

present alive in the Gardens a Wattled Lapwing (Lobivanellus

lobatus) which we have had tor nearly 20 years, and an Oyster-

eatcher which we have had for 13 years. The average durations

on the list are in most cases between 1 and 2 years. In the

period in question these birds were kept in an artificially warmed

aviary, with or without access to open air, the best figures relating

to species usually kept in an aviary with open-air cages. The

structure of the alimentary canal of these birds is on the same

general type—relatively very long and of slender calibre, with in

the less specialized forms the ceca and hind-gut relatively more

capacious, but seldom conspicuously so. The usual failure with

common European birds is noteworthy, and would appear still

more Clearly if I had not omitted from the figures numerous cases

of individuals that lived less than a month at the Gardens. The

evidence appears to show that in proportion to their size

Limicolous birds have a high potential longevity, but that either

their viability is constitutionally low, or the conditions of captivity

are unfavourable to them.

Capt. Flower’s highest record is a Lapwing still alive after

6 years’ duration.

GAVIA.

STERCORARIIDA, Tdi Gaetan ae

Stercorarius, whole genus ..... Pe nea ae 14 14-6 67

S. antarcticus (Antarctic Skua) 6 18 67

S. buffoni (Buffon’s My Fal 2 2

S. crepidaius (Richardson’s ,, ) 2 1 1

S. pomatorhinus (Pomatorhine ,, ) 5 19 AT

LARD.

Phaéthusa magnirostris (Great-billed 1 2 2

Tern)

Gelochelidon anglica (Gull-billed .,, ) 1 10 10

Sterna, whole genus ..........5.02.-.5.3. 12 116 74

S. cantiaca (Sandwich Tern) ............ 4 2°7 3

S. fluviatilis (Common ,, )............ i 16 74

S. minuta (Little SD ae ties att 1 16 16

Larus, smnole. semus) Geen eee 190 42°5 257

LL. argentatus (Herring Gull)............ 34 27°8 99

L. atricilla (Laughing ,, )............ 6 26 50

L. brunneicephalus (Brown-headed 1 55 55

Gull)

I. cachinnans (Yellow-legged Herring 11 OD 230

Gull)

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 53)

ao Tata, be faineatin, Sonat

Larus canus (Common Gull)............ 27 30°9 (Gy

L. cirrhocephalus (Ashy-headed Gull) 3 47 94

L. dominicanus (Dominican alee 10] 114

L. franklini (Franklin’s a eeo 10 16

L. fuscus (Lesser Black-backed ,, ) 11 32°8 97

L. gelastes (Arabian madi rage 10 10

L. glaucus (Glaucous are 58°5 156

L. ichthyaétus (Great Black-headed ,, ) 2 6°5 7

L. leucopterus (Iceland mado L 93 93.

L. marinus (Greater Black-backed ,, ) 8 74 187

L. nove-hollandie (Jameson's ae elt 60 255,

L. ridibundus (Black-headed ,, ) 57 AQ) 257

Pagophila eburnea (Ivory a) ea: 20 5D

Rissa tridactyla (Kittiwake) ............ 18 10°5 55

Summary of Gavie.

Mr. J. H. Gurney (loc. cit. p. 40) mentions Herring Gulls of

44 and 21 years, Lesser Black-backed Gulls of 30 and 32 years,

and a Great Skua of 24 years. The highest figures on my list are

a Black-headed and a Jameson’s Gull, each of over 21 years’

duration—and a Yellow-legged Herring Gull of over 19 years’

duration. Within the divisions there is no apparent correlation

between size and either maximum or average durations. The

Skuas and Terns lived on an average under 2 years, most of the

Gulls more than 4 years. The Gulls were kept, during the period

covered, in out-door aviaries, or if they were pinioned, in open

paddocks. The alimentary tract of all the Gavie is relatively

long and of narrow calibre: the ceca and hind-gut are very short

and uncapacious except in the Skuas, when they are of moderate

length. The group, in proportion to its size, shows only moderate

longevity and viability.

Capt. Flower’s only high record is that of a Lesser Black- backed

Gull alive after 11 years’ duration.

TUBINARES.

PurFINIDZ. No. of — Av. dur. Max. dur.

Indiv. in months. in months.

Paijenius, whole genus 2. ........c.. 0.05, 6 16 4

P. anglorum (Manx Shearwater) ...... 3 2 4

P. kuhli (Cinereous Puffin) «............ 2 1 1

P. major (Greater Shearwater) ......... 1 1 |

Fulmarus glacialis (Fulmar) ............ 4 10 38

[ do not know of any published information as to the longevity

of the Tubinares, and the fate of the few examples on my list is

only sufficient to show that they have very bad lives in captivity.

532 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

PYGOPODES.

CoLYMBIDH. ee ee aa

Tachybaptes fluviatilis (Little Grebe) 10 10°3 50

Eehmophorus major (Great Selec 9 9

Podiceps cristatus (Crested, “)" 3 4-3 Wil

P. griseigena (Bedenecked 5) eat J i

Colymbus arcticus (Black-throated 1 1 1

Diver)

C. glacialis' (Great Northern. ,,) 2 Sel 10

C. septentrionalis (Red-throated ,, ) 6 5) Ve

ALCIDA.

Allcattorda (Razor eee cee 12 9 5A

Lomvia troile (Common Guillemot) ... 33 6 52

Uria grylle (Black itn a) ee eG) 1 2

Fratercula arctica (Puffin) ............... 11 5 16

In the case of this group also, I do not know of published

information. The records I can give are very bad; of 86 examples

the average duration is under a year, and in most cases much

under a year, whilst the figures would have been still worse if I

had included a considerable number of Little Grebes, Crested

Grebes, Razorbills, and Puftins that failed to livea month. In

all these birds the intestinal tract is of moderate length, in

proportion to size; even in the Divers the ceca and hind-gut are

uncapacious, whilst in the others they are vestigial. Jam unaware

of anything in the structure of these birds from which we might

infer a constitutional lack of viability as compared with, for

instance, Gulls, and the conditions under which the attempt has

been made to keep them are similar. I think that their lack of

viability in captivity must be partly psychological, and to be

referred to the set of causes which lower the average duration

in captivity of most of the familiar mammals and birds of

Europe.

IMPENNES. No. of Av. duy. Max. dur.

Indiv. in months. in months.

Aptenodytes pennant: (King Penguin) 9 3°6 11

Pygosceles teniatus (Gentoo fe CS 2°5 7

Spheniscus, whole genus ............... 29 17-5 135

S. demersus (Black-footed Penguin)... 24 20 135

S. humboldti (Humboldt’s is sae) 2 5 9

S. magellanicus (Jackass a Vee eaace 5 8

Fudyptes, whole genus .................. 16 13°5 88

H. antipodum (Yellow - crowned 2 6°5 12

Penguin)

H. chrysocome (Rock-hopper ,, ) 6 12 58

i. pachyrhynchus(Wide-beaked ,, ) 5 4 8

E. selateri (Sclater’s Sed) sete tes 37 88

RELATIVE VIABILITY IN MAMMALS AWD BIRDS. DBs

In this group also I am unaware of published figures as to

longevity. To the maximum of over 11 years’ duration in the

case of a Black-footed Penguin, I am able to add that of another

example of the same species, now alive in the Gardens after a

duration of 123 years, whilst there are several of over 6 years.

The average duration shewn by the records is bad, in all cases

under two years, In most under one. Penguins have an

excessively long and slender alimentary tract, but Ae hind-gut and

ceca are vestigial. In the period under consideration they were

kept for the most part ina house with artificial heat and without

access to fresh air, but some of the larger forms had free access to

open air and they were by no means the most successful. I think

it must be inferred that these birds havea good potential longevity,

but that their viability in captivity, at least, is low.

No. of Ay. dur. Max. dur.

CRYPTURI. Indiv, im months. in months.

Tinamus solitarias (Solitary Tinamou). 6 30 78

Crypturus, whole genus .............., 24 12:7 66

C. noctivagus (Banded Tinamou) ... 10 13-4 AD

C. obsoletus (Obsolete _,, ) a ean Neve 4

C. tataupa = (Tataupa _s,, ant ae Al 66

C. undulatus (Undulated ,,° =) :.. 1 6 6

Rhynchotus perdicarius (Chilian 2 . 245 26

Tinamou).

R. rufescens (Rufous Tinamou) ..... 66 19°6 102

Nothura maculosa (Spotted Tinamou) 48 12 3D

In the case of Tinamous, once more I am unaware of the

existence of published records as to longevity. The maximum

duration of 84 years (in the case of a Rufous Tinamou) and the

average durations, which in most cases are under 2 years, show

that these birds are rather less viable than the Guans, with

which they may be compared in size and to some extent in habits.

They have been kept under similar conditions—in an aviary with

warmed shelters and open-air compartments. The structure of

the alimentary canal differs from that of Gallinaceous birds chiefly

in the relative shortness and want of capacity of the whole tract

andin the much greater relative capacity and length of hind-gut

and ceca.

APTERYGES. No. of _Av. dur. Max. dur.

Indiv. in months. in mouths.

Apteryx, whole genus ......2,.:2..0.0+. 22 48 240

A. australis (Kiwi) inseam ee 1 97, OF

A. haasti (Haast’s Kiw i) pie edie 2 555 4G

A, mantelli (Mantell’s ,, ) ......... 9 50°5 240)

A. owenit (Owen's al) Leen 10 38°8 ial iy

Mr. J. H. Gurney (doe. ett. p. 40) records an Australian Apteryx

which died after having lived 20 years in our Gardens; this is

Proc. Zoo. Soc.—1911, No. XXXV, 35

534 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

probably the same specimen recorded im my list as a Mantell’s

Apteryx. Although the Kiwis are small if considered in com-

parison with Struthious birds, they are actually large birds, and

the figures of maximum and average durations cannot be taken

as showing that these birds have a high viability in proportion

to their size. They are nocturnal, and have generally been con-

fined in small shelters in a heated aviary. ‘The alimentary tract

is of slender calibre, and not long in proportion to the size of

ihe bird, but the hind-gut and especially the cca are extremely

capacious.

% a if Te KS Ae al

CASUARIT.* die, ree ae

Caswarius, whole genus :.............. 12 30°8 186

C. caswarius (Common or Ceram 11 ol 110

Cassowary).

C. c. beccarii (Beccari’s Cassowary)... 7 16 24

OC, Gm Salonen (ID ANISS 5 css I 20 20

CO. c. salvadori (Salvadori’s_,, eRe ee) 22a) 61

C'. c. intensus (Blue-necked ,, | erent 17 29

C. c. violicollis (Violet-necked 6 26 38

Cassowary).

CO. c. australis (Australian - 5 64 186

C’. bicarunculatus (T'wo-wattled 5 34 86

Cassowary).

C. uniappendiculatus (One-wattled 10 1359 43

Cassowary).

C. w. occipitalis (Jobi Island One- 1 10 10

wattled Cassowary).

C. uw. aurantiacus (Kastern One- 1 8 8

wattled Cassowary).

C. wu. rufotinctus (Red-tinted One- 2 ely 26

wattled Cassowary).

C. phitipr (Selater’s 43 1 jl 31

C. papwanus (Westermann’s ,, ) 5 45 85

C. p. edwardst (Milne-Edwards’s 2 14 15

Cassowary).

C. picticoll’s (Painted-necked ,, ) 1 29 29

Ombennetia (Nloonuils)\sierern tata eee 5 78 174

Dromeus nove-hollandic (Kmu) 5 13-4 29

Mr. J. H. Gurney (loc. cit. p. 40) has recorded an Emu of 28

years, and a Westermann’s Cassowary of 26 years. The figures on

iy list are very much lower, the maximum duration being 153

years fora Cassowary. In the period considered they were kept in

a warmed house with open-air yards, but to what extent they were

allowed free access to the yards, I am unable to say. The figures

* (In the synonymy of the Cassowaries I have followed Mr. L. W. Rothschild,

Trans. Zool. Soc. vol. xv. p. 109. |

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 535

of average duration vary, but these and the maximum durations

both show that at least under thesconditions of captivity, Casso-

waries and Emus have a low viability in proportion to their

size. It is interesting to notice that the Emus, although larger

than most of the Cassowaries, appear to be less viable. In the

section on Struthiones which follows I refer to the conditions of

the alimentary tract of these birds.

SEU EELONES: tng ON Ga osteo rat cohen!

Struthio camelus (Ostrich) ............ 25 15-4 56

S. molybdophanes (Somali Ostrich)... 4 25 62

Rhea americana (Common Rhea)... 51 aod Lee 80

R. darwin (Darwin's) ">." )ee 6 9 12

R. macrorhyncha (Great-billed Rhea) ial 17-6 43

Metchnikoff (op. cit. p. 78) has collected some information as

to the longevity of Ostriches and their allies, and has come to

the conclusion that 35 years would be an extreme age to be

attained by these gigantic birds. The figures on my list show

much shorter maximum durations, whilst the figures of average

duration are still lower. It is interesting to note, that taking

Kiwis, Cassowaries, Emus, Rheas, and Ostriches together, not

only is there no direct correlation between size and average

and maximum durations, but the familiar relation is reversed.

The smallest birds, the Kiwis, according to my tables, show

the highest viability; next come the Cassowaries, whilst the

Ostriches, much the largest birds, are lowest. In the Kiwi

and Cassowary, the ceca, and hind-gut, although moderately

capacious, are not specially so in proportion to the remainder of

the alimentary tract ; whilst in the Rheas, Emus, and above all

in the Ostrich, the hind-gut and ceca are relatively enormously

long and capacious, out of all proportion to the condition in

any other group of birds and recalling the structure of herbi-

vorous mammals. In their case there is the maximum opportunity

for intestinal putrefaction, and these large Struthious birds, in

proportion to their size, appear to have the lowest viability of

all birds. Capt. Flower’s only high record is an Ostrich alive

after 12 years’ duration.

SuMMARY or AYES.

Constitutional differences in Longevity.

Putting together the information so carefully collected by

Mr. Gurney, which of course related to maximum durations and

potential longevity, and the results which I record here, we can

draw the conclusion that very real constitutional differences

in viability and longevity exist amongst the different families

of birds. The chief difficulty is to distinguish between low

viability due to special conditions of captivity and constitutional

35%

536 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

low viability. It cannot be supposed that birds which are

resident in Europe, or in England, have a lower constitutional

viability than their congeners in more distant parts of the

world, and yet, if we were to form conclusions as to the-via-

bility of such birds (or mammals) from the evidence given by

captive examples, we should have to assign to them a very low

position. So also insectivorous birds have usually a low via-

bility in captivity, but this may well be due to the difficulty of

providing them with suitable food. So also, if we were limited

to knowledge of parrots under the usual conditions of a Parrot-

house, we should have a very erroneous view of their poten-

tialities.

Passerine birds are long-lived and hardy. Almost all of them

have a potential longevity probably well over 20 years, and some

of the larger forms may reach to over 60 years. The least hardy

in captivity are Warblers, Tits, Pipits, Wagtails, Bulbuls, Orioles,

Tanagers, and Larks. Tyrants; Bell-birds, and so forth, which are

generally regarded as a lower grade of Passerines, appear less

viable and shorter-lived.

Of Picarian birds, Colies, Hornbills, Motmots, Toucans, and

Barbets have potential longevities approaching those of Passerines,

but markedly lower, whilst their viabilities are relatively good ;

Woodpeckers, Kingfishers, Hoopoes, Bee-eaters, Rollers, Frog-

mouths, Cuckoos, and Touracous have much lower potential

longevities and are less hardy. Owls have a potential longevity

certainly extending to over 50 years, but their viability appears

to be constitutionally low, their short average durations in cap-

tivity not depending altogether on any specially unfavourable

conditions.

Parrots have a high potential longevity, certainly ranging well

over 50 years. Their viability is almost certainly good, and

their low average durations must be assigned to unfavourable

conditions in captivity.

Diurnal birds of prey have potential longevities probably

exceeding those of parrots. Their viabilities are better than

those of Owls, the conditions in captivity being in neither case

satisfactory.

Steganopodes have potential longevities ranging up to about

50 years, with very good viability ; here seems no constitutional

reason to assign a low viability to Gannets and Cormorants,

and their tre atment in captivity does not differ much from that

of Pelicans. The actual figures show that they have had much

lower average and maximum durations than Pelicans, but this

most pr obably i is to be associated with the failure in the case of

many other European birds.

Herons, Storks, Spoonbills and Ibises appear to have a poten-

tial longevity of over 30 years, and a fairly good viability.

Tbises appear to have a better viability than the other members

of the group.

RELATIVE VIABILITY IN MAMMALS AND BIRDS. Doi

Flamingoes have a potential longevity certainly considerably

over 20 years, and have a high viability in captivity.

Screamers appear to have rather low potential longevity and

viability, certainly much lower than those of Flamingoes or

Geese.

Swans, Ducks, and Geese have a longevity most probably going

beyond 50 years, and appear to have a good viability.

Doves and Pigeons have a _ potential longevity certainly

reaching to about 30 years and have a very good viability.

Sand-grouse apparently are considerably lower than Pigeons

both in potential and average duration.

Gallinaceous birds have a potential longevity ranging about

20 years, and only a fair viability. Megapodes, Curassows, and

Guans seem to be most hardy and the longest livers.

The Hemipodes have considerably lower potential longevity

and viability than Gallinaceous birds of about the same size.

_ Rails, Crakes, Porphyrios, and Gallinules appear to have poten-

tial longevities of not more than about 20 years, and have

relatively low viability

Of the Alectorides, Cranes have a potential longevity almost

certainly exceeding 50 years and a very good viability.

Kagus have a still better viability and pr obably a somewhat

sunilar potential longevity. Bustards are the lowest of the group

both in longevity and viability.

Limicolous birds have a potential longevity which is high in

proportion to their size and must be set down as at least about

30 years. Hither their constitutional viability is low or the

conditions of captivity are specially unfavourable to them.

Gulls and their allies certainly have a potential longevity of at

least 30 yeas, but their viability, at least under the conditions

of captivity, is relatively low.

Puffins; Divers, Guillemots, and so forth have so poor a via-

bility in captivity that no conclusions can be drawn as to their

potential longevity.

Penguins have a potential longevity of at least over 12 years,

but their viability under the conditions of captivity, in the period

covered by my records, is low.

Tinamous appear to have a low potential longevity, certainly

not much over 12 years, and a relatively poor viability,

The potential longevity of Struthious birds is certainly under

50 years, and their viability is relatively low.

Longevity and Size.

A very summary inspection of the tables of figures I have

given will show that for birds as a whole there is no constant

relation between potential longevity and size. The correlation

does not exist even if only size and actual longevity be taken. An

Ostrich is much bigger than a Crow or a Parrot, and yet cannot

538 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

attain so great an age. Nor does the correlation exist between

size and viability, for if the same sets of birds be compared, it

will be seen that the average expectation of life is much

better in the case of the smaller birds. If, instead of absolute

correlation, relative correlation be considered, the case is more

striking. | An Ostrich must be several hundred times larger and

heavier than many birds which could outlive it, and which on

the average do outlive it. If the comparison between birds and

mammals be taken, it 1s equally clear that, apart from such a case

as that of the whale for which a very great age is assumed, and

that of man, there are many birds with potential longevities

equal to those of the longest-lived mammals. Moreover, if the

two. Classes be compared from the point of view of longevity in

proportion to size, the result is still more striking. On the

whole, group by group, mammals are much larger than birds, her-

bivorous mammals than herbivorous birds, frugivorous mammals

than frugivorous birds, omnivorous or carnivorous mammals than

omnivorous or carnivorous birds. And yet, group by group, if the

figures be compared, it becomes more and more obvious that birds

approach and often surpass mammals in longevity and viability.

Metchnikoff has already called attention to this difference and

has associated it with the anatomical fact that the most striking

difterence between the alimentary tracts of mammals and birds

is in the greater relative length, complexity and capacity of the

hind-gut in the former group. The facts that mammals with

relatively reduced and uncapacious hind-guts and ceca (such as

the Carnivora) tend to have longer lives m proportion to their

size, and that birds like the Struthious birds, which have relatively

long and capacious hind-guts and ceca, have short lives in

proportion to their size, still further confirm and support

Metchnikoff’s general proposition.

Within certain groups of birds the correlation between size and

potential longevity appears to exist. It can be traced with more

or less clearness in the case of Hagles, Owls, Passeres, and Picarian

birds. But even in this limited fashion it is far from being

universal. Probably Swans have a higher longevity than Ducks

and Geese, but I cannot find any similar differences if Geese and

Ducks be compared. Kagus appear to live as long as Cranes, Ibises

better than Storks or Herons: there is no trace of the correlation

in. the case of Rails, or Pigeons or Gulls, and amongst Struthious

birds the conditions are apparently reversed, and the Kiwi is

hardier than the Ostrich.

I should hesitate to say that the differences in the constitutional

longevities of birds could be associated universally with- the

structure of the alimentary canal, although there are some indi-

cations pointing im that ditcenions The Struthious birds and the

Screamers have large and capacious hind-guts and exca, and in

proportion to their size have very short lives. Sand-grouse

difter from Pigeons in having capacious ceca and have relatively

BLATIVE VIABILITY IN MAMMALS AND BIRDS. 539

shorter lives. Owls have large ceca, Hagles and their allies have

vestigial ceca, and Owls in proportion to their size appear to

have lower potential and average longevities.

Many large birds such as Eagles, Vultures, Pelicans, Flamin-

goes, and Cranes, can attain great ages, but similar ages can be

attained by much smaller birds such as Parrots and Crows. In

proportion ‘to their size, it is quite clear that Passeres have the

longest potential and average durations; and that Parrots,

Pigeons, and Limicolous bir ds, similarly in propoition to size,

come next, There are few generalizations more difficult to

support with convincing argument than those which relate to the

relative degrees of specialization to be assigned to natural groups

of animals, but I think that many ornithologists would agree with

me if I were to say that Passeres, Parrots, Pigeons, and Limicolous

birds were relatively high types, standing far away from whatever

we may imagine the primitive bird-type to-have been. And I

think that the most certain general conclusion which I may

draw regarding the relative longey ity and viability of the groups

of birds is, that in propor tion to size, longevity and viability

increase with perfection of organization. The further a bird has

advanced along the lines of evolution of the bird group, the more

viable it is. No doubt shortening of the hind-gut, reduction of

the number of eggs in a clutch, increased care of the young, may

all be taken as stages in the perfection of the bird-type, and so

have a general association with increased longevity, apart from any

direct influence which any one of them may have

Viability, Temperature, and Open Air.

I cannot see that there is any relation between the climate

from which birds come and their viability in captivity. Nor

would such a relation be expected on general grounds, for birds,

by their covering of feathers and the perfection of their vascular

and respiratory systems, are well adapted to resist exposure to

changes in temperature. If food be abundant, and if there be

protection from draught, they seem remarkably indifferent to the

temperature of the air. The interpretation, however, of their

viabilities is obscured by special factors which existed in the period

with which the figures deal, and which exist in most menageries.

The comparison that is to be desired is a direct contrast between

the viabilities of birds kept in the interior of a warmed house, and

birds kept with little or no artificial heat and free exposure to the

open air. The numbers of species and of individuals usually kept

in captivity is much greater in the case of birds than in the case

of mammals, and from the smaller size of birds and their more

gvegarious habits, it is usually possible to keep numbers of

individuals of the same or of different species in the same aviary.

The most familiar contrast in the conditions under which birds

are kept, is that between one or two individuals in a small cage in

540 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

a warmed house, and considerable numbers of individuals in an

aviary or enclosure giving room for exercise, and access to the open

air. Now although many birds can be kept together, and are kept

together, many casualties occur. A good deal of fighting and

bullying takes place, and in practice, it is extraordinarily difficult

to secure that every bird in the collection gets its due share of

food, and is not worried by its messmates. Moreover, there is

special danger of individuals which are affected by a communi-

cvble disease, or by parasites, infecting their healthy neighbours.

Birds in separate cages avoid these evils; they are safe from

bullying, they are properly fed, and if they are out of condition

temporarily, have a chance of recovery. If we discount the very

considerable advantage of protection that the Parrots obtained

from their housing in separate cages, it will be seen from com-

parison of their average durations with their known high specific

longevity, how poorly the device of keeping them in a heated

house has answered. In a majority of cases, birds that have had

free access to open air have lived in communities, and before

judging of the advantage of fresh air, we have to discount the

pevils arising from their communal life. With these limitations

and corrections, | think it is as clear an inference in the case of

birds as in that of mammals, that making protection from

temperature more important than the access to fresh air has

failed in practice.

GENERAL SUMMARY.

(1) This memo brings together the records of the duration of

life in captivity of over 20,000 individual mammals and birds, and

contains information which, so far as I can ascertain, has never

before been set out systematically. The individual facts are

imperfect, inasmuch as the sources did not record the age, or

condition, of the individuals at the time of their reception in the

Collection. The probable effect of these imperfections is to lower

the figures both of averageand of maximum duration. I hope to

have shewn that from the point of view of exact knowledge and

for practical reasons, it is important that those in charge of living

animals should take steps to record such, information as exactly

as possible. Every individual should be observed for some time

after its arrival, and the date when it is passed into the general

collection as apparently in good condition should be recorded,

whilst, if the exact age be unknown, it should be classified as

infantile, adult, or aged.. The houses, enclosures, or general con-

ditions under which it is kept, with the date and nature of any

changes of these, should be recorded. Finally the date, cause of

death, and any further notes as to age should be entered. With

such a system, in course of time, very exact conclusions could

be obtamed as to the viability and longevity of different kinds

of animals, and as to the effect of the modes of keeping them in

captivity.

(2) Following Ray Lankester, I distinguish between potential

RELATIVE VIABILILY IN MAMMALS AND BIRDS 5Al

longevity *, the limit of age which an individual could attain in

the most favourable circumstances, and average or specific

longevity, the average age to which the animals of a species

attain under the natural conditions to which the species has

become adapted. The difference between these two, I suggest is

a measure of the severity of the conditions to which the species

is subjected. In the case of the vast majority of animals it is

impossible to get information as to either average or potential

longevity. What information we have, is derived chiefly from

knowledge of animals in captivity, and in this memoir I have been

able to set down further information about species which have

already been discussed by J. H. Gurney, Metchnikoff, Brehm and

others, and to add information regarding many species about

which nothing has been recorded hitherto.

(3) I apply the principle of comparison between average and

potential longevities to the case of animals in captivity, by

discussing the meaning of average duration and maximum

duration. I suggest that a measure of the effect of the conditions

to which animals are subjected in captivity may be obtained by

comparison of the maximum duration with what may be known

from other sources of the potential longevity, and by comparison

of the average duration with the maximum duration. If the

individual facts were collected in such a fashion as I have

suggested in paragraph 1 of this general. summary, I think the

new method would lead to incontrovertible results, but even with

the facts at my disposal, conclusions of fair validity can be drawn.

It is necessary to note, however, that the figures of individual

duration could be used to reveal more, if they were plotted out in

curves instead of being used to give an arithmetical] mean. The

grouping of the individual cases with regard to the mean is of

great practical importance. In some cases, for instance, I have

pointed out that the majority of the cases were grouped towards

the two extremes, that the individuals for the most part either

* [Lankester (¢.c. p.27) pointed out that some organisms, such as fish, molluses,

large crustacea, sea-corals, and many trees, appear to have no fixed potential longevity

but to persist until they perish from disease, or are overwhelmed by some accident,

whilst others, such as man, appear to havea set period to the possible duration of their

lives. he mammals and birds with which I am concerned in this memoir belong

to the second category. But I doubt if there be a fundamental distinction between

the categories. First,as Lankester stated, the individuality of an organism like a tree,

or a colonial invertebrate, differs from that of most organisms, and must be separately

considered from the point of view of potential longevity. Next, certain organisms,

such as many fish, appear to have more indefinite limits of growth (possibly related

to mechanical factors, of similar importance in the case of aquatic mammals such

as the Cetacea) than the majority of animals, the adult dimensions of which vary

within narrow limits; definite or indefinite potential longevity may be related with

definite or indefinite growth. (See also H. Spencer, ‘ Principles of Biology,’ revised

edition, 1898, vol. i. p. 135). I am: more inclined to suppose that potential longevity

varies with structure in the widest sense, including under the term structure the

consequences of wear and tear, and auto-poisoning from the slow accumulation of

waste-products. All pieces of machinery, from a motor-car to a cork-screw, have

a potential longevity, dependent in the main on their structure, and apart from the

quality of their manufacture, varying fairly closely with their complexity. Although

our attention is arrested by extreme cases, these are only terms in a series. |

542 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

died in the first few months after arrival (by birth or otherwise)

or survived to an age approaching the maximum, whilst in other

cases there was a mortality rate steadily increasing with the

length of residence in captivity. Obviously, the different curves

that would be displayed by different animals, if the individual

cases were plotted out in such a fashion, would give valuable

information as to the special periods of danger in the case of

different animals, and inform us if there were cases where the

mortality was the result of progressive and cumulative effects of

captivity. As, however, the facts at my disposal did not include

any information as to the age or condition of the animals on

reception, I came to the conclusion that it would be a waste of

time to plot out curves.

(4) Unfortunately, collections of animals have to be considered

from the financial point of view. The money value of an animal

to a collection, that is to say, the price which can be judiciously

paid for it, depends on its rarity, its attractiveness to the popular

and scientific clients of the institution, and on the length of time

it may be expected to live. A gorilla, for instance, is rarer and

much more attractive than a chimpanzee, but as a gorilla has

a very much worse expectation of life in captivity than a chim-

panzee, its money value may be smaller. The facts that I have

brought together form, so far as I know, the first beginning of

a practical guide to the value of animals considered from this point

of view.

(5) I have been able to bring together a large series of facts

with regard to the relation between size and longevity. Taken in

the broadest way, it is true that large animals may be expected

to live longer than small animals, and in the case of very closely

allied creatures the relation is frequently close. This may be

associated partly with the effects of the accumulation of waste

products. The cubical capacity of an animal increases much more

rapidly than its linear dimensions, and of two animals of similar

structure and constitution, the larger may take longer to be

poisoned by its own waste products. Of course many simpler

factors are involved—such as the greater resistance of a more

bulky animal to rapid changes in the temperature of the air or

water in which it lives. On the other hand, difference in

longevity is not in exact proportion to size, and otlfer con-

stitutional factors are more important. Birds, for instance, in

proportion to size, have higher potential longevities than mammals;

whilst within the classes, orders, and even families, there are many

eases where difference in size is overborne by other constitutional

differences. It seems to be the case that, in proportion to their

size, the more highly developed members of a group are able to

live longer than their lower kin. There are also a very large

number of cases, where increased viability and longevity are

associated, as Metchnikoff suggested, with a relatively low capacity

of the hind-gut.

(6) It has long been known to aviculturists that common

RELATIVE VIABILITY IN MAMMALS AND BIRCS. 543

British birds are often very difficult to keep in captivity. I have

shown that this difficulty occurs in so large a number of different

kinds of British birds and mammals, that it may be laid down as

a curious principle (to which naturally some exceptions exist) that

British birds and mammals have a lower viability in captivity in

England than their immediate allies from any other part of the

world. This remarkable circumstance is in the first place, in the

strict sense of the word, accidental, and finds partial explanation

in a cause independent of the constitution of the animals. Most

captured examples of wild species either are weakly individuals,

or are injured or frightened by the process of capture: a heavy

mortality is to be expected. Tn the case of animals that come from

a distance, much of the heavy mortality takes place before arrival

or, because of the additional evil effects of the conditions during

transit, and the survivors which reach their destination are rela-

tively strong and hardy. On the other hand, local animals

reach their destination in a shorter time, and the heavy mortality

takes place inside the Gardens. But this explanation is not wholly

sufficient to cover the cases, and I think it may be assumed that

wild birds and mammals in Britain have acquired an intolerance

of man, without which, unfortunately, they would not have been

allowed to maintain their existence. This psychological acquisition

presses heavily on them in captivity. It appears to be the case

that an opposite process of selection is taking place in the parks

of great cities, and that wild birds in particular are learning not

to fear man. It would be interesting to know if wild birds

taken in a London park lived better in captivity than birds of

the same species from country districts.

(7) The climate from which a bird or mammal comes has the

smallest possible relation to its viability in captivity. A. Heilprin

(Distribution of Animals, Int. Sci. Series, vol. lvii., 1887, p. 35)

pointed out the error of the common belief as to climate being

the principal factor that regulates or controls the distribu-

tion of animals. Amongst mammals and birds a vast majority

of species anfl genera regarded as tropical have an actual or

recent range into temperate or even frigid climates. Of those

now limited to the tropics, still fewer are accustomed to a steady

temperature. Some range periodically or occasionally to altitudes

where great cold occurs; others, by exposure to the intense

radiation of the dry air of plains at night, regularly endure cold

going down to freezing-point ; whilst many inhabitants of tropical

forests (which we naturally associate with steamy heat) must be

subjected to great cold in their nocturnal wanderings on the

summits of tall trees. The abundance of thick hair and fur and

of close feathering amongst tropical creatures is a clear indication

that their lifeis not spent basking in tropical sunlight. Mammals

and birds have the power of maintaining their internal temperature

at a normal that varies only within an extremely narrow range,

notwithstanding the temperature changes in their environment ;

and I do not doubt, not only that they can endure considerable

544 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

cold, but that even rapid and considerable changes of temperature

are a necessary stimulus to their viability.

On the other hand, there are cases where the change to the

climate of London is certainly trying, but these occur not amongst

tropical but temperate or arctic animals. The production of a thick

coat against the onset of winter cold, and its dotting when summer

approaches, appear not to be direct reactions to temperature but

organic rhythms adapted to the seasons in the natural habitat of

the animals. Thusanimals from the far North, accustomed to the

short arctic summer, retain their thick coats im this climate long

after their possession is cumbersome. Still worse is the case of

animals brought from South temperate zones, which begin to

moult their thick coats when our winter, their summer, approaches.

I have not information as to how soon if ever these rhythms

readjust themselves to the reversed conditions. It is clear, on

the other hand, that part at least of the mechanism by which

coats become warmer, 1s direct stimulation from the surrounding

temperature, and such different creatures as Carnivora and

Baboons rapidly get better coats when exposed to the open air.

‘The seasonal rhythms connected with breeding are also a cause

of mortality to young or adults, when the conditions of climate

are reversed. Birds from the Southern hemisphere, if they

survive, appear to readjust themselves in this respect; it is more

doubtful if mammals do so.

(8) The idea that it is a fundamental] necessity to protect neal Wes

adult mammals and birds from cold by providing them with

artificial heat is fallacious. The supreme necessity is free access

to open air. In most cases this should be combined with shelter

from rain and wind, and in some cases the shelter should be

supplied with artificial heat—perhaps often even in excess of what

is now customary—but only so far as it can be arranged without

any detriment to fresh air. I think this is probably specially

important in the case of nocturnal animals; as we are accustomed

to see these asleep all day in the warmest corner given them, we

are disposed to forget that at night they move about actively often

in great cold.

For all mammals and birds stexdy exposure to an even tem-

perature is unnatural and unhealthy; change is a necessary

stimulus, and permanent existence indoors is the wom possible

condition for viability and longevity.*

* [When this memoir was read my friend Dr. Leiper called my attention to the

undoubted fact that the problem of the duration of life of animals in captivity is com-

plicated by the eftects of parasites. In the course of my paper (see pp. 470 and 540)

I have made reference to this subject, pointing out that some of the evil caused by

confinement in warmed houses is undoubtedly not the direct result of absence of

tresh air, or of artificial heat, but comes about mediately inasmuch as the conditions

are specially favourable to parasites. It is true of course that any argument as to

the potential longevity of animals based on maximum durations in captivity is’

subject to the effects of parasitism. J hope that the work of Mr. Plimmer, Mr. Beddard,

Dr. Leiper, Dr. Nicoll, and Mr. Coventry, who are now systematically examining

the parasites at the Gardens, will before long enable us to eliminate some at least of

these evils. |

RELATIVE VIABILITY IN MAMMALS AND BIRDS. 5AD5

(9) The tabulation and analysis of the records in this memoir

has been a laborious task which has not been lightened by the

knowledge that the materials were imperfect. But I trust that

it may be a step towards obtaining systematic and more exact

knowledge on the subject. Such knowledge i is the only sure basis

for that improvement in the condition of animals in captivity

which those in control of Zoological Collections desire to bring

about. The peculiarity by which a menagerie is distinguished

from a museum is that its zoological specimens are alive. Not

the obtaining of rare animals, nor the addition to the records of

* species new. to the Collection ” should be the chief glory of a

Zoological Society, but that. Collection should be judged most

valuable and successful in which the average duration of life of its

inhabitants approaches potential duration most closely.

Nore on THE THEory or LONGEVITY.

With three notable exceptions, those of Lankester, Weis-

mann, and Metchnikoff, discussions of the theory of longevity

may be dismissed briefly here, partly beeause they have been

treated fully in the literature of the subject *, and partly

because they do not pretend to explain why the gift of life has

been measured out to different animals in unequal portions.

Bacon made a careful summary of the known facts, but refrained

from general conclusions. ‘‘ In tame creatures” he wrote, as cited

by Lankester from Basil Montagu’s translation, “ their degenerate

life corrupteth them, in wild creatures their exposing to all

weathers often intercepteth them; neither do those things which

may seem concomitants give any furtherance to this information

(the greatness of their bodies, their time of bearing in the womb,

the number of their young ones, the time of their growth, and

the rest), in regard that these things are intermixed, and some-

times they concur, sometimes they sever.” Buffon thought that

there was a relation between the total duration of life and the

period of growth, and set down the ratio between the two as 6 or

7 to 1. Flour ens followed Buffon’s idea, but took as the limit of

erowth, the age at which the long bones unite with their

epiphyses, estimating that the ratio of the longevity to the period

of growth was as 5 to 1. Bunge, without calculating an exact

ratio, pointed out that there was a frequent relation between the

longevity and the time taken by a new born animal to double its

weight. Such calculations at the best are limited in their

application to the higher animals, and even amongst these have

to encounter many exceptions : so far as they go “they must be

taken as secondary and comparatively accidental correlations.

Ray Lankester’s early contribution (¢. c. p. 71) was a serious

* See specially E. Ray. Lankester, ¢. c.; KE. Metchnikof!, ¢. c. p. 39, and article

Longevity, im the XI. edition of the Eneyclopedia Britannica.

546 DR. P. CHALMERS MITCHELL ON LONGEVITY AND

addition to knowledge. Having distinguished between potential

and average specific longevities, and shown that the latter were

determined to a large extent by accidents outside the constitution

of the species (destruction by enemies, diseases and so forth), he

attributed the former to constitutional causes of which the most

important were the degrees of evolution or individuation and the

amounts of personal and generative expenditure. High individua-

tion, and low expenditure were to be associated with great potential

longevity.

A. Weismann, in his famous essay Ueber das Dawer des Lebens

{translated in Essays upon Heredity, Oxford 1889), examined the

various constitutional explanations of the duration of life. He

admitted that small animals might be expected to run through

the cycle of life more rapidly than large animals; that if the

period of growth were long it might expand the total duration

of life; that creatures in which the metabolic processes were

extremely active, might finish their career more quickly (as Lotze

suggested in his Microcosmus) than slow-living creatures; that

the rates of personal and of reproductive expenditure had some

influence. But he urged that the application of these various

principles was only partial and led to so many inconsistencies -

that no constant correlation could be established. He came to

the conclusion that duration of life was really dependent upon

adaptation to external conditions, that its length was governed

by the needs of the species and was regulated by the same process

as that by which the structure and the other functions of an

organism were adapted to the environment.

In adaptation to the environment, Weismann urged, it is the

prosperity of the species and not that of the individual that is

concerned. That species is most successful which contains at any

time the largest number of vigorous adults, and as every organism

in the vicissitudes of life becomes to a certain extent worn and

dilapidated, it is not to the advantage of the species that indi-

viduals should live too long. As soon as the business of repro-

duction has been successfully accomplished, the advantage of an

individual to the species is gone, and the sooner it disappears the

better. Animals are in fact wound up to go for the requisite

time,and no longer. The mechanism Weismann suggested was a

limitation of the possible number of cell generations, admitting

that this was hypothetical. The requisite length of time was

determined by the reproductive habits of the animals. Slow

breeding, for instance the production of a small number at a

birth or in a season, long brood-care either embryonic or post-

embryonic, and all the various circumstances by which an organism

or pair of organisms require longer time to replace themselves

by their younger and fresher children, stretched out the dura-

tion of life; whilst rapid reproduction, absence of brood-care and

so forth, contracted it. By the process of natural selection the

incidence of death was adapted to the needs of the species.

RELATIVE VIAPILITY IN MAMMALS AND BIRDS. SAT

Obviously where there was a heavy incidence of death on the

young, a longer life would be required to secure replacement of

the parent by successfully reared young.

Weismann (é. ¢. p. 11) brought together a very remarkable and

cogent series of cases showing that a constant correlation existed

between the reproductive powers and the duration of life of the

individuals of a species, and his theory has an extremely im-

portant place in the history of the subject. It is to be noted,

however, that although he appreciated the probability that a very

large number of deaths were due to external causes (enemies,

diseases, accidents, untoward conditions) he did not explicitly

work out the important relation to his theory of Lankester’s

distinction between average specific longevity and potential

longevity. It is clear that if the duration of life be fixed by some

internal cause such as the limitation of cell-reproduction, it must

be the potential longevity and not the average specific longevity

that is affected; but if the majority of animals perish from

accidents from without, it is difficult to see how their potential

longevity could be influenced by selection.

Metchnikoff’s most interesting work has still further increased

the diticulty in the way of accepting Weismann’s theory.

Metchnikoff investigated the causes of death in a very large

number of cases, including those of insects which die very soon

after having pac their eggs, and came to the conclusion that it is

extremely doubtful if natural death occurs except in the rarest

possible cases. Most animals perish long before they have reached

an age at all approaching what may be regarded as their potential

longevity. In the cases that survive the accidents of youth or early

maturity, another series of accidents take effect. The changes of

senility are induced from without; the various degenerations

are brought into existence by the absorption into the system of

various kinds of poisons, and these whether they are due to the

exudations of the microbes of intestinal putrefaction, or to those

of special diseases, are entirely external to the constitution.

Senile animals perish because they can no longer resist common

diseases or common accidents. It would be only in a world

where the most perfect hygiene reigned, and from which the

bacteria and microbes of diseases had been dri ‘iven, that the

majority of the members of a species would attain the potential

age, and that death would come on them from purely constitu-

tional causes. And so the conception of the duration of animal

life being an adaptation to breeding habits, produced by the

operation of selection, disappears.

I cannot doubt but that the average specific longevity is

the dominating factor in animal life. At every stage of their

existence animals are assaulted from without by enemies, diseases,

and their whole environment. To say that is merely to restate

the existence of the struggle for existence, the fundamental

proposition of the theory of natural selection. If favourable

548 LONGEVITY AND RELATIVE VIABILITY IN MAMMALS AND BIRDS.

variations arise, that is to say changes in diathesis, structure,

function or habit, which enable the possessors of these to make a

better fight against enemies, or diseases, or any part of their hostile

environment, then we expect such variations to be preserved, and,

consequently, the constitution of the organism to be modified in

the direction of adaptation to its environment. It is no part of

my present argument to discuss whether such variations appear in

response to the environment, or by large or small stages; these

are wide qnestions not germane to the present issue. But they

do arise, and in sucha fashion there have come about increases or

decreases in size, advances in structure or degenerations, protective

colours and weapons—the whole armoury offered by variation to

the choice of natural selection. Such improved powers of resistance

to the environment may increase the average specific longevity,

and indirectly may affect the potential longevity. But the attain-

ment of the latter is too rare an event to come under the direct

control of natural selection.

There is no more important part of the structure and function

of an organism than that associated with reproduction. The

impulses and instincts connected with fertilization, the time,

duration and frequency of rutting, the numbers of ova that

are fertilized and allowed to develope, the duration of embryonic

development, the amount and character of post-natal brood-care,

the whole apparatus and machinery of the reproductive process,

present an extraordinary series of modifications from species to

species, and from individual to individual. They are subject

to much variation amongst individuals at the present time,

and it may well be presumed that they have offered a rich .

material to natural selection in the past. Weismann has

established a correlation between longevity and reproduction,

but I think his interpretation of the correlation must be reversed.

Tt is not longevity that has become adapted to reproduction, but

the rate of reproduction has been adapted not to potential

longevity, but to average specific longevity. Average specific

longevity is simply a measure of the pressure of conditions on a

species, and its constitution, including its mode of reproduction,

has been shaped in conformity with the pressure of these

conditions. Potential longevity is a bye-product of the process ;

it depends directly on the constitution, and is simply the result of

the fact that if the constitution be adapted to meet the average

mortality, it must be able to outlive, and will outlive, the average

duration of life in a number of fortunate cases.

ON NEW PARASITIC NEMATODES FROM TROPICAL AFRICA, 549

25. Some new Parasitic Nematodes from Tropical Africa.

By Rosert T. Lerreer, D.Sc., M.B., F.Z.S.

[Received April 4, 1911: Read April 25, 1911.]

(Text-figures 140-144.)

In the following paper I give a brief description of a number

of new genera that have recently come under my notice in the

course of an examination of helminthic material collected by me

during a visit to Kast Africa, Uganda and the Soudan, whilst a

member of the Egyptian Government Survey in 1907, and of

further material sent to me at the London School of Tropical

Medicine by members of the Colonial Medical Service.

NEMATODA.

Family ANCYLOSTOMID 4,

Genus ACHEILOSTOMA, gen. n.

Species SIMPSONI, sp. n. (‘ype sp.)

Host: a large rodent. Locality: Nigeria.

A large number of specimens was collected from the ali-

mentary canal and preserved in 70 °/, alcohol. The males measure

17 mm. in length, the females 23 mm. in length. The former

are less numerous than the latter. The cuticle shows marked

transverse striation in the anterior end of the body. The indi-

vidual striz are so deeply cut as to give the edge of the cuticle

in this region a serrated appearance. Ata distance of 0:8 mm.

from the anterior end are two large stout lateral papille curving

backwards and shaped like large thorns (text-fig. 140, A, /.p. p.550).

The anterior end of the body is bent dorsally only very slightly,

showing that the oral aperture, which is surrounded by a thick

collar, 0-004 mm. deep, is almost but not quite terminal. The

mouth capsule shows similar characters to those found in other

members of this family. There is a large median ventral tooth

and two fan-shaped inner teeth guarding the cesophageal entrance

to the buccal capsule dorsally (text-fig. 140,D). The buceal capsule

has a chitinous wall, and resembles that of Vecator rather than that

of Ancylostomum. 'The most striking feature of the genus, how-

ever, is the entire absence of teeth or cutting-plates guarding the

entrance to the oral aperture. There is no indication either of a

corona radiata; so that although the buccal cavity of the parasite

recalls that of the Ancylostomes in its internal characters, the oral

aperture reminds one rather of the Bunostomes.

The csophagus measures 0°95 mm. in length and shows no

special characters. In the female the genital pore lies almost

at the centre of the body. The eggs in the uterus measure

0-055 mm. x 0°032 mm. and usually contain a few segments.

In the male, the posterior end of the body is surrounded by a

Proc. Zoot. Soc.—1911, No. XXXVI. 36

D5) DR. ROBERT T. LEIPER ON NEW

well developed bursa that projects much more ventrally than

dorsally (text-fig. 140, B). The genital cone (text-fig. 140, B, g.c.)

is greatly developed and can be seen projecting between the two

sides of the bursa and the dorsalinembrane. The dorsal membrane

is distinctly delimited from the lateral halves of the bursa and is

very much shorter (text-fig. 140, C).

Text-fig. 140.

Acheilostoma simpsoni.

A. Anterior extremity showing mouth capsule m.c., wsophagus @s., Jateral

cervical papilla /.p.

B. Bursa of male, lateral view: g.c., genital cone; sp., spicules.

C. Bursa showing division of dorsal ray.

D. Base of mouth capsule showing chitinous folds of the wall of the capsule

guarding the cesophageal orifice.

The dorsal ray is short and thick and bifurcates about halfway

along its length to form two stout rays, which are again split into

two subdivisions as they reach the edge of the bursal membrane.

The externo-dorsal rays leave the dorsal ray from about the

middle of the thick undivided portion. The ventral rays are

fairly well developed and are united to their tips. All the lateral

rays, middle and posterior, run to the bursal edge almost side by

side, but the anterior ray, which has a thickness almost equal to

the two central rays combined, takes a separate course, ending

about one-third of the distance from the midlateral to the

ventral rays. (Collected by J. J. Simpson, Esq.)

fe |

—

PARASITIC NEMATODES FROM TROPICAL AFRICA. 5

Family STRONGY LID &.

Subfamily CyLrcosromiIn %.

Genus CYLINDROPHARYNX, gen. n.

Species BREVICAUDA, sp. n. (Type sp.)

Host: Zebra. Locality: British Hast Africa.

The Zebra is the subject of a considerable number of parasites

that occur normally in the Horse, but in addition it appears to

have found a few peculiar to itself. While examining a number

of Cylicostominz cleared in glycerine, I noticed a number of

specimens that appeared to have an abnormally deep buccal

capsule. The shape of this capsule struck me at once as of generic

value, and on further search I was soon able to distinguish in the

material two distinct species showing this special character.

To the naked eye appearances the species of this genus resemble

exactly those of Cylicostomwm. The mouth is surrounded by a

collar bearing a double corona radiata. In C. brevicauda, the

type species, the buccal capsule is cylindrical, and has a thick

chitinous wall measuring 0-4 mm. in depth and 0-1 mm. in

transverse diameter. The wsophagus is exceedingly short and

fleshy, measuring 0-46 min. in length and 0°18 mm. in greatest

diameter. The nerve-ring surrounds the esophagus immediately

behind its union with the buccal capsule (text-fig. 141, A, p. 552).

Tn the female, the posterior end tapers very quickly from a diameter

of 0°25 mm. at the level of the vulva to a pencil-like point. The

vulva opens at 0°7 mm., the anus at 0°18 mm. from the extremity

(text-fig. 141, C). In the male the bursa shows a characteristic

disposition of the rays. The posterior ray is split to its base into

two portions, and each of these is again split as far into an internal

and external ray. The external ray is bifurcated as it reaches the

edge of the bursal membrane (text-fig. 141, D). The genital cone

is very large and protuberant. The spicules are long and filiform,

measuring 1mm.in length. There isa stout thick uneven accessory

piece 0-2 mm. in length (text-fig. 141, B). The edge of the bursal

membrane is finely serrated as in the genus Triodontophorus.

CYLINDROPHARYNX LONGICAUDA, Sp. Nn.

Host: Zebra. Locality: British East Africa.

This species closely resembles C. brevicauda, but the buccal

capsule is smaller, measuring only 0°23 x 0-07 mm. The cesophagus

is more slender, measuring 0°42 x 0:12 mm. (text-fig. 142, A).

In the female the tail is long and pointed. The vulva is situated

much farther forward in this species, opening at 1°55 mm., whilst

the anus opens 0°32 mm. from the posterior extremity (text-

fig. 142, D). In the male the spicules are shorter, 07 mm. The

accessory piece is less developed and shorter, 0°13 mm. The genital

cone is more globular (text-fig. 142, B). The rays differ more par-

ticularly in that the external branch of the posterior is undivided

(text-fig. 142,C). (Collected by Dr. J. T. C. Johnson.)

DR. ROBERT T, LEIPER ON NEW

Sil

Text-fig. 141.

Cylindropharynx brevicauda.

A. Anterior extremity showing deep cylindrical pharynx (ph.) and short muscular

cesophagus (cs.), nerve-ring (nv.), and chyle imtestine (c.in.).

B. Bursa of male showing the long spicules (sp.’, sp.’’) and well-developed accessory

piece (acc.). The externo-dorsal ray is bifurcated at its tip. ‘There is a

large protruding genital cone (g.c.).

C. Posterior extremity of female showing relationship of vulva (v.) and anus (an.)

to tip of tail.

D. Dorsal rays of bursa as seen from ventral surface; the edge of the bursa is

serrated.

Subfamily STRONGYLINA.

Genus TRACHYPHARYNX, gen. n.

Species NIGERIA, sp. n. (Type sp.)

Host: a large rodent. Locality: Nigeria.

Among the above specimens were a few shorter and more

stunipy forms that on microscopical examination proved to belong

to quite a different family. The specimens were few in number

and badly preserved. The diagnosis is based upon the anterior

end of the body, mainly because in both sexes the posterior end

was obscured and embedded in a thick brown prostatic secretion.

The males measure 11 mm. in length and the females 15 mm. in

length, and 0-46 mm.in breadth. They taper a little at either end.

PARASITIC NEMATODES FROM TROPICAL AFRICA, 553

Text-fig. 142.

Cylindropharyne longicauda.

Explanation of individual figures as in text-fig. 141.

The oral aperture is quite terminal, and is surrounded by a well

developed corona radiata highly reminiscent of that found in the

species Strongylus equinus. The number of leaves in the external

crown could not be determined exactly. The internal crown

was distinctly visible as a series of comma-like projections,

apparently corresponding in number to those of the external

erown. The oral capsule is a wide pear-shaped cavity, the

anterior third of which is surrounded by a thick wall of

cuticle, the posterior two-thirds being almost entirely replaced by

a layer of cesophageal musculature, which is however covered by

a thin layer of cuticle bearing a large number of round cuticular

knobs that project into the buccal cavity. The esophagus mea-

sures 1°25 mm. from the base of the cuticular portion of the

buccal capsule to its termination in the chyle intestine. Where

554 DR. ROBERT T. LEIPER ON NEW

the cesophageal musculature supports and encloses the buccal

cavity the muscular tissue is only 0:006 mm. in thickness, but in

the posterior third the cesophageal musculature attains a thickness

of 0°13 mm,

Text-fig. 143.

Trachypharynex nigeria.

Anterior extremity showing buccal capsule.

The females were full of eggs, and although the details of

the posterior end of the body could not be made out exactly, the

vulva appeared to be situated a short distance in front of the

anus, which in turn was situated almost at the posterior end of

the body. (Collected by J. J. Simpson, Esq.)

Family SPIROPTERIDS.

SPIROPTERA UGANDA, sp. n.

Host: Monkey. Locality: Uganda.

In the intestine of a Monkey a fragment of a male worm was

found. The anterior portion, having been cut off in the process

of opening the gut, was unfortunately lost. The essential points

are, however, shown in text-fig. 144, which illustrates the

posterior end, showing the papille and other specific characters.

The papille are large, and arranged as four pairs of pedunculated

preanals and five pairs of postanals, three of these being pedun-

PARASITIC NEMATODES FROM TROPICAL AFRICA. 555

culated, the terminal two pairs being sessile. On either side, the

cuticle is expanded to form an alar ledge about 0°2 mm. in breadth.

The expansions are marked on the ventral aspect by a longitudinal

Text-fig. 144.

Spiroptera uganda.

Posterior extremity of male: al., ale; sp., spicules.

striation similar to that commonly found in other species of the

genus Spiroptera. The spicules are remarkably short and almost

equal, measuring 0°2 and 0°3 mm. in length respectively.

ADDENDUM.

CoppoLDINA nom. nov.

* for Cobboldia Leiper 1910, preoce.

_ In my memoir on the “ Entozoa of the Hippopotamus” pub-

lished in the Proceedings of the Zoological Society for 1910,

at page 255, I described and figured a new genus Oobboldia

having as type Cobboldia vivipara, sp. n.

Mr. Hassall and Dr. Schouteden have kindly drawn my

attention to the fact that Cobboldia has been used already for

a genus of Insects and is therefore not available. I propose

to replace Cobboldia Leiper 1910, by the new generic name

Cobboldina.

R. T. Lerper, D.Se., F.Z.8.

May 1911.

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No. 93.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.

March 21st, 1911.

Dr, S. F. Harmer, M.A., F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The SECRETARY read a Report on the additions that had been

made to the Society’s Menagerie during the month of February

Mr. R. I. Pocock, F.Z.S8., Superintendent of the Gardens,

exhibited :—

(1) A pair of Otter cubs about seven weeks old, which were

found under a landing-stage at Tewkesbury, and presented to the

Society by Mr. W. Baring Bingham, F.Z.8.

(2) A specimen of the North American Black-footed Polecat

(Putorius nigripes), recently received in exchange from the

Zoological Society of Washington, a species furnishing, in

Mr. Pocock’s opinion, an admirable instance of “ warning

coloration.”

(3) The skin of a Chacma Baboon (Papio porcarius), from

Potchefstroom in the Transvaal, representing an apparently un-

described subspecies, for which the name griseipes was proposed,

in allusion to the speckled coloration of the hands and feet, a

character in which it differed markedly from the typical black-

handed Chacmas of Cape Colony.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of St# Shillings per annum, payable in advance,

(4) The frontlet and antlers of a specimen of the Manchurian

Wapiti (Cervus xanthopygus), to show variation in the structure

of the two antlers, one having a short “ bez”-tine nearly midway

between the ‘“ brow” and the “ trez,” the other a long “ bez ”-tine

rising close to the “ brow ”-tine.

Mr. D. Ssru-Smita, F.Z.8., the Society’s Curator of Birds,

exhibited a living Hybrid Duck, which was believed to be a cross

between the White-eyed Pochard (A ythya nyroca) and the Marbled

Duck (Marmaronetia angustirostris). It was hatched at Scampston

Hall, Yorkshire, from a clutch of eggs laid by a Marbled Duck

at Lilford Hall, Northamptonshire. The bird displayed the

characteristic markings of Marmaronetta, but the general colour

was dark reddish brown. It had been presented to the Society

by Mr. W. H. St. Quintin, F.Z.5.

Mrs. E. W. Srxton presented a paper, communicated by Dr. W.

T. Cauman, F.ZS8., “On the Amphipod Genus Leptocheirus.”

In preparing this revision of the genus the type specimens of

nearly all the species had been examined. The author found

that in some cases different stages of growth had been described

by different authors as distinct species, in other cases the in-

adequacy of the original descriptions and figures had Jed to the

introduction of a number of unnecessary synonyms. ‘The speci-

mens of Zaddach’s Z. pilosus and Grube’s LZ. guttatus had been

examined and re-described, and the number of valid species in

the genus was now seven, namely: ZL. pilosus, L. pinguis,

L. hirsutimanus, L. pectinatus, L. guttatus, L. aberrans, and

LL. bispinosus.

Messrs. J. Lewis Bonnotz, M.A., F.L.S., F.Z.8S., and F. W.

SmautEy, F.Z.S., read their paper on Inheritance of Colour in

Pigeons, which dealt with the first results of a long series of

experiments. Although chiefly Mendelian in character, the

authors laid stress on the fact that in several respects their results

seemed to point to a further law or laws, which were able to

modify the expected Mendelian results and in regard to which

the Mendelian theory offered no satisfactory solution. They

showed, for instance, that in Chequer and Blue matings there

was a regular tendency to an over-production of Chequers, in

the Grizzle matings the tendency was to an over-production of

Blues. Another point apparently inexplicable on the Mendelian

hypothesis was the difference in shades of the same colour; by

disregarding these and considering them all as gametically iden-

tical, the results gave approximately the expected proportions,

but, on the other hand, the shades of the different birds un-

doubtedly affected their progeny, and hence the gametes must

also have been affected. The preponderance of a certain sex in

a particular colour was also noted, as well as an increase of white

in successive generations. No explanation of these phenomena

was put forward, as further experiments were still in progress.

In dealing with the purely Mendelian aspect of the results

the following points were clearly brought forward :—(1) Silver is

dilute Blue. (2) Blue is dominant to Silver. (3) Chequering

and Grizzling are both dominant to absence of pattern. (4) Griz-

zling is dominant to Chequering. (5) A Mealy is a Grizzled bird

with the White replaced entirely or partially by Red. (6) Red in

a Mealy is dominant to White, hence a Mealy is dominant to a

Grizzle. (7) White and Grizzling combine to have a common

inheritance. (8) Red combines with Grizzling in the same way

as White.

Dr. G. StewaRDson Brapy, M.D., F.R.S., C.M.Z.S., presented

a paper on Marine Ostracoda from Madeira, based on specimens

collected by the Rev. Canon Norman, F.R.S8., in the spring of

1897. Apart from the species described as new to science, the

collection was interesting as extending the known range of several

species from the European and North Atlantic areas much farther

southward, though not quite into the tropical zone.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, April 4th, 1911, at half-past Hight

oclock p.M., when the following communications will be

made :—

ier: Res Lererr, FZ:

Demonstration of Nematode Parasites obtained from Animals

in the Gardens.

2. F. E. Bepparp, M.A., F.R.S., F.Z.8.

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.—No. I. On some Mammalian Tapeworms.

2. J. A. Morcu.

On the Natural History of Whalebone Whales.

The following communications have been received :—

1. Witi1am Nicott, M.A., D.8c., M.B., ¥.Z:8.

On Three new Trematodes from Reptiles.

2. Dr. R. W. Suureipt, C.M.Z.8.

On the Comparative Osteology of Cercopithecus and Callithrix

(C. sabeus and CO. callitrichus and C. jacchus).

Communications intended for the Scientific Meetings should

be addressed to ;

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Socrery oF Lonpon,

Recent’s Park, Lonpon, N.W.

March 28th, 1911.

No. 94.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

April 4th, 1911.

Dr, Henry Woopwarp, F.R.S., Vice-President,

in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

Dr. H. B. Fantuam, F.Z.8., and Miss AnnrzE Porter, D.Sc.,

exhibited some diseased bees and combs infected with a minute

pathogenic Protozoal parasite, apparently the same as Vosema

apis found by Zander and Doflein in diseased bees in Bavaria.

Microscopic preparations and drawings of the parasite, Vosema

apis, were also shown, as well as healthy bees and combs in

contrast. The material exhibited was obtained from Cambridge-

shire and Hertfordshire in March 1911. The infected combs

were brown in colour instead of the normal yellow, while the

infected bees suffered from a sort of dry dysentery which rapidly

proved fatal. The pathogenic agent of this dry dysentery,

Nosema apis, formed thousands of minute spores which fouled

the hive, while infection was probably spread to new hives by

hungry, weakly bees attempting to enter healthy hives. The

spores, about 2 to 3 by 4 to 6, were the resistant and cross-

infective stages of the Protozodn. The parasite Nosema apis

was closely allied to that of pébrine, the silkworm disease due to

Nosema bombycis. The trophozoite and pansporoblast stages of

NV. apis had been observed, as well as some spores with polar

filaments extruded. Like WV. bombycis, the bee-parasite was

possibly capable of hereditary infection, as infected bee-larve had

been found. The only certain destructive agent of the Micro-

sporidian spores was fire, and all infected bees and hives, and any

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Str Shillings per annum, payable in advance.

débris therefrom should be most carefully burned. In the opinion

of the exhibitors, the Microsporidian parasite, Vosema apis, had

been responsible for much of the bee-disease recorded in this

country since 1906, especially in 1906, 1907, and 1911. The

exhibitors first noticed the parasite in 1906 in diseased bees ob-

tained from the Isle of Wight; its full significance was grasped

in 1907, but owing to the difficulty of obtaining material the

exhibitors’ results were not published. As much attention was

now being directed to “‘bee-disease,” the exhibitors briefly recorded

their observations. It was not asseited that microsporidiosis was

the only disease of bees current in Great Britain at present, as

Dr. Malden had investigated a bacillary infection in bees. Micro-

sporidiosis had probably been introduced from the Continent into

British apiaries.

Dr. R. T. Luiprr, F.Z.S., gave a demonstration of Nematode

parasites obtained from animals in the Zoological Gardens during

the year ending November 1910.

The collection contained a number of new forms, of which a

systematic account will be published later. Among the more

interesting of the known forms were fictularia plagiostoma

from a Palm-Civet, a number of species of Polydelphis from

various Pythons, Dicheilomena horrida from the South American

Ostrich, and Dictyocaulus jilaria from the lungs of Sheep.

It was noticed that whereas intestinal parasites were almost

wholly collected from animals that had not lived in the Gardens

for more than six months, those of which the normal habitat and

food were the internal tissues of the host occurred in animals

that had been confined in the Gardens for several years. Thus,

an undescribed /laria was found in a Lemur after four years, and

filaria australis ina Wallaby after two and a half years’ captivity.

In all these cases the number of parasites obtained was small,

and could have had little or no effect upon the heaith of the

host. There was a remarkable preponderance of female forms.

From these observations it appeared that the change of food

and general conditions obtaining in the Gardens were unfavourable

to the continued existence of the intestinal parasites an animal

may harbour on its admission. The number of cases of auto-

and re-infection during captivity was strikingly small, and bore

testimony to the cleanly surroundings in which the animals were

kept. In four cases only was there evidence of the occurrence

of accumulative infection in the Gardens :—

1, A number of Giant Toads died from lung infection with

Lhabdias bufonis.

2. The Wolves appeared to be heavily infected with Ascaris

canis,

3. A Sheep died from pneumonie condition resulting from

an intense infection with Dictyocaulus filaria.

4, The Tortoises had Oxyuriasis.

In all these cases repeated infection undoubtedly had followed

from contamination of food and drink with feces containing

eggs of the parasite. The infection could be eliminated by steam

sterilisation of the cages, or still more easily by changing the

species of animal living in the particular paddocks or cages, for

Helminthes were often peculiarly selective as regards their hosts,

and those flourishing in one animal sometimes found it impossible

to continue their life even in closely allied forms.

Sir E. Ray Lanxester, K.C.B., F.R.S., F.Z.8., exhibited a

special Supplement of the ‘Field’ newspaper dealing with the

British non-migratory Trout, and called attention to this new

medium for the publication of scientific observations requiring

ulustration.

The Secretary read a letter from the Governor of Seychelles,

received through the Secretary of State for the Colonies, on the

subject of the herd of Land-Tortoises at Government House,

Mahé.

Mr. R. I. Pocock, F.LS., F.Z.8., Superintendent of the

Gardens, exhibited the body of a newly-born cub of the Masked

Palm-Civet (Paradoxurus larvatus) from Szechuen, and after

pointing out its differences from the adult, drew attention to a

peculiar abnormality in the left foreleg.

Mr. F. E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the

Society, read a paper on some Mammalian Tapeworms which

had been collected from animals that had died in the Society’s

Gardens. This collection was the result of nearly two years’

examination of a very large number of animals, but did not

contain a very large number of species. Tapeworms were

by no means so common as other parasitic worms, particularly

Nematodes, which were the most abundant among the animals

in the Gardens.

Dr. 8. F. Harmer, M.A., F.R.S., V.P.Z.S., communicated a

paper by Mr. J. A. Morch, of Christiania, on the Natural History

of Whalebone Whales, drawing attention to, and throwing light

upon, some of the problems connected with the migrations of the

larger Cetacea.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, April 25th, 1911, at half-past Hight

o'clock -p.m., when the following communications will be

made :—

1. Mr. D. Seru-Suiru, F.Z.8.

Exhibition of Lantern-slides :—

(a) Penguins in Moult.

(6) Wild Swainson’s Lorikeets.

2. Dr. R. T. Lerper, F.Z.S.

Some new Parasitic Nematodes from Tropical Africa.

3. Dr. Curusert Curisty, F.Z.S.

On a Collection of Antelope and other Skins from the

Chagwe Forests, Uganda.

4, Wiuttam Nicout, M.A., D.Sec., M.B., F.Z.S.

On Three new Trematodes from Reptiles.

The following paper has been received :—

H. W. Marert-Tims, M.D., F.Z.8.

Tooth-Germs in a Kangaroo.

Communications intended for the Scientific Meetings should

be addressed to

P, CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,

ReceEnt’s Parx, Lonpon, N.W.

April 11th, 1911.

No. 95.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

April 25th, 1911.

Dr. S. F. Harmer, M.A., F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The SecRETARY read a Report on the additions that had been

made to the Society’s Menagerie during the month of March

USS

Mr. D. Seru-Suirn, F.Z.8. the Society’s Curator of Birds,

exhibited :—

(1) A nest of the Grey Struthidea or Apostle Bird (Struthidea

cinerea), composed entirely of mud, and built on a branch in the

Western Aviary.

(2) Lantern-slides from photographs of the King Penguin

(Aptenodytes pennanti) and Black-footed Penguins (Spheniscus

demersus) showing the method of moulting.

(3) Lantern-slides from photographs of a number of wild

Swainson’s Lorikeets (Zrichoglossus nove-hollandice), kindly sent

by Mrs. Innes, of Mackay, North Queensland. These birds came

in large numbers to feed daily at a table, on syrup provided,

settling without fear upon the head, shoulders, and arms of the

lady who fed them.

My. C. Tate Recan, M.A., F.Z.S8., exhibited a series of lantern-

slides of scales of the Salmon (Salmo salar), and showed how the

life-history of the fish could be read from its scales.

* This Abstract is published by the Society at ils offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Sia Shillings per annum, payable in advance.

Dr. Witi1am Nicout, M.A., F.Z.S., exhibited some preparations

from a Hare which showed an interesting and unique patho-

logical condition. The liver was extensively invaded with

calcareous nodules which were due to a parasitic Nematode of

the family Z7ichotrachelide. 'The worms were so coiled up that

they could not be extricated entire, but the ova, which were

present in enormous numbers, showed that they must belong to

a peculiar genus. The structure of the eggs was remarkable in

displaying a double-layered shell, with a markedly papillated

surface. No similar condition had been hitherto recorded from

the Hare or Rabbit, but an analogous condition had been met

with fairly frequently in Rats. It was still impossible to say

whether the conditions in the two animals were caused by the

same species of parasite, but experiments with a view to deter-

mining this were in progress. ‘The Hare further showed a large

infection with Trichostrongylus retorteformis, and a slight, purely

intestinal, infection with Coccidium cuniculi. There were also

signs of recent parturition and of an inflammatory condition of

the uterus. Whether this or the liver disease was the cause

of death, which took place under remarkable circumstances, is

doubtful.

Dr. Curupert Curisty, F.Z.8., exhibited part of a collection

of skins of mammals and reptiles obtained by him in Uganda,

which included those of the Antelope, Leopard, Civet, Hyrax, &c.,

and drew attention to a rare form of Dendrohyrax, D. emini,

and to the skin of a melanistic form of the Civet.

Dr. Wiuuiam Nicoutt, M.A., F.Z.8., read a paper on Three

new Trematodes from Reptiles, from material received from the

Society’s Prosectorium. The specimens were interesting as

forming an important addition to our knowledge of the large

variety of forms which inhabited the air-passages and anterior

coil of the alimentary canal of reptiles and batrachians.

Dr. R. T. Leper, M.B., F.Z.S., read a paper on some Parasitic

Nematodes from Tropical Africa, and gave a brief description of

a number of new genera. The paper was based on helminthic

material he had collected during a visit to Hast Africa, Uganda,

and the Soudan in 1907, and on material sent to him by members

of the Colonial Medical Service.

Mr. OuprieLD THomas, F.R.S., F.Z.8., read a paper, the

fourteenth of the series, on Mammals collected in Southern

Shen-si, Central China, by Mr. Malcolm Anderson, for the Duke

of Bedford’s Exploration of Hastern Asia. ‘The region explored

was in the Great Pe-ling (or Tsin-ling) range, that divides

Northern from Southern China, many of the specimens coming

from the sacred mountain Tai-pei-san, where several of the most

interesting forms were ebtained.

Of these by far the most striking was a new species of Takin

(Budorcas), readily distinguishable by its uniform golden buffy

colour from the Sze-chuen species (B. tibefanus). In the adult

of this fine animal the coloration was wholly buffy, the darkening

of the ears, dorsal line, hinder back and limbs found in B. tibe-

tamus being absent, and there was scarcely a trace even of the

dark facial patch so prominent in that animal. The new species

was proposed to be called Budorcas bedfordi, and female no. 2190

was selected as the type.

Other new forms were as follows :—

ARCTONYX LEUCOLEMUS ORESTES, subsp. n.

Lighter than in true lewcolemus. Dark orbital patch reduced

to a narrow rim round eye.

Head and body 570 mm.; tail 195; skull 132.

Hab. Tsin-ling Mts., alt. 12,000’. Type. Female. No. 2191.

Microtus CALAMORUM SUPERUS, subsp. n.

Rather larger and with decidedly longer tail than true

calamorum.

Head and body 130 mm.; tail 63; hind foot 24; skull 33.

Hab. 8. Shen-si. Type. Male. No. 2163.

OCHOTONA SYRINX, sp. n.

Near O. cansa, but larger. Bulle smaller,

Head and body 142 mm.; hind foot 28°5; skull 36:5.

Hab. Tai-pei-san. Type. Male. No. 2170.

In all, the collection contained 160 specimens referable to

30 species.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, May 9th, 1911, at half-past Hight

o'clock p.m., when the following communications will be

made :—

1. RK. 1. Pocock, F.L:S., F:Z.S.

On the Palatability of some British Insects. (Hxperiments

made in the Society’s Gardens with Arthropods (chiefly In-

sects) and Molluscs, and Notes on the significance of Mimetic

resemblances.)

9. Prof. Gitpert C. Bourneg, D.Sc., F.R.S., F.Z.S.

Contributions to the Morphology of the Group Neritoidea of

Aspidobranch Gastropods.—Part II. The Helicinide.

3. J. J. Laster, M.A., F.B.S., F.Z.S.

On the Distribution in the Pacific of the Avian Family

Megapodide.

The following papers have been received :—

1. H. W. Marert Tims, M.D., F.Z.S., and A, HopewEeLL Smiru,

L.R.0.P.

Tooth-Germs in a Kangaroo.

2. R. Broom, M.D., D.Sc., C.M.Z.S.

On the Structure of the Skull in Cynodont Reptiles.

3. The Rev. A. Mites Moss, M.A., F.Z.8., F.ELS.

On the Sphingide of Peru. With a Preface by Karu

JORDAN, Ph.D.

4, C, KE. Hetimayver, M.B.O.U.

A Contribution to the Ornithology of Western Colombia.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL SocrETY OF LONDON,

Regent's Park, Lonpon, N.W.

May 2nd, 1911.

No. 96.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.”

May 9th, 1911.

EK. G. B. MEapre-Wa po, Esq., Vice-President,

in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

Mr. R. I. Pococr, F.R.S., F.L.8., Superintendent of the Gardens,

exhibited some of the hair of the “ puppy coat” of a Grey Seal

(Halicherus grypus), which was caught at Barmouth, in Merioneth,

at the end of April. When received at the Gardens at the

beginning of May this Seal was covered, with exception of the

head and flippers, with longish woolly white hair, the last of

which was moulted on May 7th. Most authorities state that

Grey Seals are born in the autumn, not later than about the

middle of October, and that the puppy coat is shed from a month

to six weeks later. Allowing six weeks for the retention of its

puppy coat, this Welsh Grey Seal must have been born near the

middle of March, a date in tolerably close agreement with the

date, namely the end of February, given by Cneiff for the birth

of these Seals in the Gulf of Bothnia. It is, therefore, quite clear

that these Seals breed both in the early spring and the autumn.

Mr. E. G. BouLenceEr exhibited some living male specimens of

the Midwife Toad (Alytes obstetricans) carrying the eggs. He

also exhibited a number of the detached eggs to show the manner

in which they were strung together.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Siv Shillings per annum, payable in advance.

Mr, A. E. Anpersow exhibited a large number of photographs

of the more important fossil mammals in the Department of

Vertebrate Paleontology of the American Museum of Natural

History, New York, showing the methods of mounting fossil

skeletons. For comparison, a set of photographs was exhibited

with the skeleton supports eliminated from view, thus adding to

the pictorial value of pose in the specimens.

Dr. P. Cuatmers Mircuety, F.R.S., Secretary of the Society,

gave an account, in the absence of the author, of a Memoir by

Lt.-Col. Nrvinne Manpmrs, R.A.M.C., F.Z.8., on the phenomena

of Mimicry amongst Butterflies in Bourbon, Mauritius, and Ceylon.

The author had investigated the habits by observation and ex-

periment of the insectivorous reptiles and birds of these islands,

and had been unable to accept the view that their relations to

butterflies were such as to be effective in producing Batesian or

Miillerian mimicry.

Mr. BR. I. Pococx, F.R.S., F.L.S., read a paper on the Palata-

bility of some British Insects, with Notes on the significance of

Mimetic resemblances, and said that at Prof. Poulton’s request

he had undertaken in the summers of 1909 and 1910 to make a

series of experiments in the Gardens to test the edibility of various

British insects, most of which were sent to him, together with

some slugs, by Dr. G. B. Longstaff. The insects comprised Lepi-

doptera, Coleoptera, Orthoptera, Hemiptera, Diptera, and Hy-

menoptera ; and the most interesting of the experiments were

those made with the Bumble-Bee (Bombus) and its mimetic fly

(Volucella bombylans) to test the theory of mimicry. The Bombus

proved to be unpalatable to nearly all birds. The birds would

try them a varying number of times. When they had learnt

their distastefulness by experience they refused to touch them,

and then when offered the Volwcella refused that likewise. A

considerable number of species of insectivorous birds were tested

in this way, and always with the same result; and the one speci-

men of Volucella bombylans that did duty for some thirty or forty

experiments went through the ordeal untouched.

Prof. G. C. Bourne, M.A., D.Sc., F.R.S., read the second

portion of his paper on the Morphology of the group Neritoidea

of the Aspidobranch Gastropods, which dealt with the Helicinide.

He stated that this family was capable, by some unknown means,

of wide dispersal across seas and oceans, and that the conditions

most suitable to its existence were found in proximity to the sea.

Jn describing the anatomy the genus Alcadia was taken as the

type, and the differences between it and the other genera were

pointed out, but the species and even the genera of Helicinide

were closely similar, anatomically, from whatever part of the

world they came.

Mr. J. J. Lister, M.A., F.R.S., F.Z.S., presented a paper

entitled “On the Distribution in the Pacific of the Avian Family

Megapodide.”

The next Meeting of the Society for Scientific Business will

be held on Tuesday, May 23rd, 1911, at half-past Hight

o'clock p.m., when the following communications will be

made :—

1. J. Stuart THomson, Ph.D.

The Aleyonaria of the Cape of Good Hope and Natal.

2. H. W. Marerr Tims, M.D., F.Z.8., and A. HopEeweL. Smits,

1G Ore.

Tooth-Germs in a Kangaroo,

3. R. Broom, M.D., D.Sc., C.M.Z.S.

On the Structure of the Skull in Cynodont Reptiles.

4, he Rey. A. Mizzs Moss, M.A., R.ZS8, RES.

On the Sphingide of Peru. With a Preface by Karu

JORDAN, Ph.D.

The following papers have been received :—

1. C. KE. Henimayr.

A Contribution to the Ornithology of Western Colombia,

2. The Hon. Paut A. Mrtuuen, F.Z.8.

On an Amphipod from the Transvaal.

3. R. LyDEKKER.

The Somali Rhinoceros and the Nigerian Klipspringer.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Socinry or Lonpon,

Rrcent’s Park, Lonpon, N.W.

May 16th, 1911,

No. 97.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

May 23rd, 1911.

Dr. A. Smita Woopwarp, F.R.S., Vice-President,

in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The Sucrerary read a Report on the Additions that had been

made to the Society's Menagerie during the month of April

ig)

Prof. ArtHuR Denby, D.Sc., F.R.S., F.Z.S., communicated a

paper by Dr. J. Stuart THomson on the Alcyonaria of the Cape

of Good Hope and Natal. The author dealt exclusively with the

order Gorgonacea, and recorded nineteen species, of which six

were described as new.

A paper entitled ‘“‘Tooth-Germs in the Wallaby (d/acropus

billardieri),” was presented by Dr. A. Hoprwexu Suiru, M.R.C.S.,

L.R.C.P., and Dr. H. W. Marert Tims, M.A., F.Z.S., F.L.S.

The material upon which their observations were based had been

kindly sent to the authors by Mr. Brooke Nicholls, of Melbourne.

It consisted of three embryos of Muacropus billardieri. The

smallest specimen (allowing for the difference in size of the adults

of different species) was considerably younger than that of any

other Diprotodont previously examined. In the upper jaw they

had identified six incisors, thus confirming M. F. Woodward’s

original statement. The functional incisors of the adult appeared

to be the 2nd, 4th, and 6th of the series, There were four

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance.

34 -

premolars, of which the Ist, 3rd, and 4th persisted. There was

also one molar tooth.

In the lower jaw, owing to the difficulty of interpreting the

conditions, it was not certain whether there were representatives

of five or six teeth in front of the premolars. Presuming there

were five, the large functional incisor of the adult was the 4th of

the series. As in the upper jaw, there were four premolars and

and one molar, the 2nd premolar not fully developing. There

were evidences of vestigial predecessors to the large lower incisor

and to pm’.

The following points of histological interest were noted :—

1. The heaping up of the ‘epithelium along the alveolar

margins, a character often supposed to be peculiar to the

Ungulates.

2. The precocious development of the enamel.

3. The compactness of the stellate reticulum of the enamel-

organ.

4. The abundant evidence of blood-vessels within the enamel-

organ, thus confirming the observations of Poulton and

Howes in the Rodents. The opposite opinion is usually

held.

5. Some slight evidence in support of the fusion of enamel-

organs. Such fusion has been recorded in the fishes and

reptiles, but not hitherto in mammals.

The Rev. A. Mites Moss, M.A., F.Z.S., F.E.S., gave a short

account of his memoir on the Sphingide of Peru, based on studies

of Lepidoptera, with special reference to the larvee, which he had

made during a three years’ residence at Lima. Dr. Karl Jordan

had assisted him with the working out of his collections, and had

described the following new species of Sphingidee :—

PROTOPARCE MOSSI Jord., sp. N.

Similar to P. seata cesiri Blanch. (1854). Both wings narrower

and more pointed. Fore wing without black streaks on the disc.

Antenna thinner. Genitalia quite different, approaching those

of P. hannibal Cr. (1779).

This paper will be published in the ‘Transactions’ in due

course.

Prof. J. P. Hitz, D.Se., communicated a paper by Dr. R

Broom, C.M.Z.8., on the Structure of the Skull in Cynodont

Reptiles. The author, after a study of all the available material

contained in the British and South African Museums, gave a

detailed comparative account, illustrated by a series of figures, of

the morphology of the skull in the chief genera of the Cyno-

dontia, including Bauria, Nythosaurus, Cynognathus, Trirachodon,

Gomphognathus, Diademodon, Sesamodon, and Melinodon. He

also discussed in some detail certain peculiarities of the Mam-

malian skull, apparently derived from a Cynodont ancestor.

— ee Se

r. C. W. Anprews, F.R.S., F.Z.8., read a paper “On a New

cs of Dinotherium from British E t st Africa.” The specimens

described were sent to the British Museum by Mr. C. W. Hoble Y,

Commissioner of Mines for British East Africa. They included

portions of the mandible with teeth, a caleaneum, and a patella

of a small species of Dinotheriwm nearly allied to D. ewvieri, from

the Lower and perhaps Middle Miocene beds of France. The

new species, which he proposed to call Dinotherium hobleyi,

differed from D. cuvieri in several particulars—e. g., the inner

anterior column of pm 3 was more distinctly developed, and the

talon of m, had a distinct tubercle on its inner side. Remains

of Rhinoceros, a giant Tortoise, Zrionyx, and Cro-odiles also

occurred. The bones were well preserved in a tough clay, and

further collecting would no doubt yield important results.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, June 13th, 1911, at half-past Hight

o'clock p.mM., when the following communications will be

made :—

eS of = Tos Tee and Lthynchotragus

from Somaliland.

The Hon. Paut A. Mernuen, F.Z.S.

On an Amphipod from the Transvaal.

3. R. LyDEKKER.

The Somali Rhinoceros and the Nigerian iXlipspringer.

4, C. BE. Henemaynr.

A Contribution to the Ornithology of Western Colombia.

5. Prof. ANGEL CABRERA, C.M.Z.S.

The Subspecies of the Spanish Ibex.

The following papers have been received :—

1. Miss Ruta Harrison.

Some Madreporaria from the Persian Gulf. With Notes on

the Memoir and some Further Notes on Pyrophyllia inflata by

Sypnry J. Hickson, M.A., D.Sc., F.R.S., F.Z8.

2. Cuarues L. Boutencer, M.A., F.Z.S.

‘On Variation in the Medusa of Merisia lyonsi.

3. Frank E. BEDDARD, M.A, E.BS., E.ZS.

Contributions to the Anatomy a Systematic Arrangement

of the Cestoidea.—No. 2. On Two New Genera of Basia dles

from Mammals.

4. R. Broom, D.Sc., C.M.ZS8.

On some new South African Permian Reptiles.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL SocreTy or Lonpoy,

Regent’s Park, Lonpon, N.W.

May 30th, 1911.

patie a yi ee

Papers (continued).

\ . Page

15. Report on the Deaths which occurred in the Zoological Gardens during 1910. By

H. G. Pruoar, F.R.S., F.Z.8., Pres.R.M.S., Pathologist to the Society

16, A Contribution to the Study of the Variations of the Spotted Salamander (Salamandra

maculosa). By Epwarp G. Boutunenr. (Plate XV. and Text-figs. 99-102.)

17. On the Mountain Nyala, Tragelaphus buxtoni. By R. Lypuxker, (Plate XVI. and

Arse LOE: Jin shearer cial elec otraats weitin dare sacha Sosusavatd sane eee aerate 348

18, Observations on different Gibbons of the Genus Hydobates now or recently living in the

Society’s Gardens, and on Symphalangus syndactylus, with Notes on Skins in the

Natural History Museum, N. Kensington. By Dr. F. D. Wutcn, F.Z.S'. ........+++5 353

19. Some New Siphonaptera from China. By Karu Jorpan, Ph.D., F.E.S., and the

Hon, N. Cuarues Roruscuinp, M.A,, F.Z.8., F.E.S8. (Text-figs. 104-124.) .......... 365

20. Contributions to the Anatomy of the Anura. By Franx H, Bzpparp, M.A., E.R.S.,

F.Z.S., Prosector to the Society. (Text-fies. 125-133.)..............cc0ecececeeeee 393

21. On the Spermatophores in Earthworms of the Genus Pheretima (=Pericheta), By

Frank EH. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the Society. (Text-figs. 184-136.) 412

22, A Rare Beaked Whale. By R. Lypexxer. (Text-figs. 187-139.) ...............0.. 420

ae Age-Phasesof the, Rorqual. By R. UypwRKaR 6. ee eta ee be eet s Sa cuee ets 423

24. On Longevity and Relative Viability in Mammals and Birds; with a Note on the

Theory of Longevity. By P. Cuatmurs Mircnunn, M.A., D.Sc., LL.D., F.R.S.,

SECie ralnya CONDOMS OCLOLYmaious tary tien. fperenersa sce tte stm stake tee amet he ere cieitea. ais! cc cueter sa teleey siete oe 425

25. Some new Parasitic Nematodes from Tropical Africa. By Roszrt T, Lerrer, D.Sc.,

ERE a Seen (Nex tefivan DAU IA4, rane acs Ah o's. vee vowels aera beeen 540

ADDENDUM.

A Correction to Dr. R. T, Lerpzr’s Memoir on the Entozoa of the Hippopotamus ........ 555

PESTER HOR Ni har ia'es ate (oleve «' Sie, cin Sys ohe ra aegis rms Reb ae | MER INRaR CO ERA tinct t SaG i

List of Council and Officers .................. 2 a ar ae eI HEME Os ol Soc li

MintvOts@ OMhenisere trast reer arerasesls, < heehee ders ote. oe Se EE EERO Ac Ont ili

ANE Rien DES Ot Coammllamnons” Ae co nloas ou Ob Hla mp Re oOaOeO DoUGUon a dsae 0000 an ix

TERE GME TENA ENE BGS AiGak Oo SERB OO CISSRELE Ed ea IRa Ene cee iar eR AIC r sae ria coe GD xvii

METS aaa eee te TC SMM Pee AN arc iA ae sh iieia’ sta stve oboe) (leis one bv Ole, #10 ah nahn fal a ane ea VE

"New Generic Terms Bene vere e tenes been eee ence eee e ete te teee whey ene ee gece XXiii

TICES Soe aig sich lie SiS SS nh eee ue ee ne AEE SORE en ein Ga eh ee ng Site oS gi

ee 243 UA

LIST OF PLATES.

1911, Parr II. (pp. 181-555).

Plate Page

II. | )

III.

IV.

v.| |

Gas Glands of Teleostel "f. 62. jen. 0 9. de see ee oie aoa oe.

VI. |

VII. |

VIII. a

ey) :

X. 1-6. Corycella brevis. 7-11. C. curta ......6. 2. se ee eee ees

XI. 1-6. Corycella curta. 7. C. brevis. 8, 9. Corycella sp. 6.

10. C. carinata: 11, 12. Coryceus gracilicaudatus .....-

XII. 1-38. Coryceus catus. 4-6. C. obtusus. 7. C. dubtus. ‘282

BOO MEMS EPMA HAR Meni tei nal dale dois rta eta tyetel cate totale |

XIII. 1-6. Coryceus murrayt. 7-9. C. andrewst .....-1..2+--++++- |

KIV. 1-4. Coryceus andrewsi, 5-9. C. dubius ..-.--++ 1.1. ..0.s 5 ee )

XV. Salamandra maculosa var. teniata, var. mollert, and ferns synaaa 323

XVI. The Mountain Nyala, Tragelaphus buxtont ...........+...-.. 348

NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively,

so that the complete reference is now P. Z. 8.1911, p.... The Distribution

is as follows :—

Part 1 issued in March.

pari oe ease June.

19 3 ” September.

Went bea Ghd December.

‘ Proceedings,’ 1911, Part I. (pp. 1-180), were published on

March 22nd, 1911.

‘The Abstracts of the Proceedings,’ Nos. 93-97,

are contained in this Part,

e918 Cl